Historical Records, Origins, and Development of the Edible Cultivar Groups of Cucurbita pepo (Cucurbitaceae) 1

Historical Records, Origins, and Development of the Edible Cultivar Groups of Cucurbita pepo (Cucurbitaceae) 1 HARRY S. PARIS 2 Cucurbita pepo includes eight groups of edible cultivars--pumpkins, scallops, acorns, crooknecks, straightnecks, vegetable marrows, cocozelles, and zucchinis. The pumpkins, scallops, and possibly the crooknecks are ancient groups that developed along separate lines of domestication in North America. Incipient forms of all the other five groups appeared in botanical herbals of Europe by 1700. Modern forms of the vegetable marrows, cocozelles, zucchinis, and acorns appeared in Europe prior to 1860, and the modern straightnecks appeared in North America by 1896. Divergence among the various groups appears to be increasing under continuing domestication due to the different demands made on cultivars grown for their immature fruits as opposed to cultivars grown for their mature fruits. An association exists between length to width ratio and stage of culinary use of the fruits in Cucurbita pepo. Cucurbita pepo L. (Cucurbitaceae) fruits occur in a myriad of shapes, sizes, and colors. Hundreds or perhaps several thousand different named cultivars are grown. Eight groups of cultivars differing distinctly in the shape of their fruits are grown for culinary purposes. These groups are the pumpkins, scallops, acorns, crooknecks, straightnecks, vegetable marrows, cocozelles, and zucchinis (Paris 1986b). The pumpkins and acorns are grown for consumption of their mature (40 d or more past anthesis) fruits. The others are grown for consumption of their immature (generally first week past anthesis) fruits. Representative immature and mature fruits of each group are depicted in Fig. 1-16. The fruits of other C. pepo cultivars are not palatable and are grown purely for decorative purposes. These are known collectively as ornamental gourds, and they occur in a wealth of shapes and colors (Bailey 1937). Some of the ornamental gourd cultivars, such as 'Flat', 'Miniature Ball', and 'Pear', differ little from wild and feral C. pepo in their phenotypic characteristics, including small fruit size and striped fruit color pattern. Cucurbita pepo, one of five cultivated species of Cucurbita, is a native of North America (Trumbull 1876; Whitaker 1947) and has been cultivated there for several thousand years (Cutler and Whitaker 1961). The cultivated forms are domesticates of wild forms from northeastern Mexico (Andres 1987) and Texas (Bailey 1943; Erwin 1938). The cultivated, edible forms of Cucurbita generally differ from wild forms by having larger and fewer seeds and fruits, non-bitterness and less fibrous nature of the fruit flesh, larger plant parts, fewer runners, and less durable and more varicolored rinds (Whitaker and Bemis 1964). Whitaker (1960), Whitaker and Bemis (1975), and Herklots (1986) suggested that the first use by early Americans of Received 4 February 1987; accepted 30 January 1988. 2 Department of Vegetable Crops, Agricultural Research Organization, Newe Ya'ar Experiment Station, P.O. Haifa, Israel. Economic Botany, 43(4), 1989, pp. 423-443 9 1989, by the New York Botanical Garden, Bronx, NY 10458

424 ECONOMIC BOTANY [VOL. 43 Fig. 1-8. Representative immature (2-5 d past anthesis) fruits of the eight Cucurbita pepo cultivar groups grown for culinary purposes. Peduncle end at top, blossom end at bottom. Fig. 1. Pumpkin 'Howden's Field'. Fig. 2. Scallop 'Patty Green Tint'. Fig. 3. Acorn 'Table King'. Fig. 4. Crookneck 'Sundance'. Fig. 5. Straightneck 'Early Prolific Straightneck'. Fig. 6. Vegetable Marrow 'Beirut'. Fig. 7. Cocozelle 'Cocozelle'. Fig. 8. Zucchini 'Bareqet'. Not to scale.

1989] PARIS: CUCURBITA PEPO 425 Cucurbita species as food was the consumption of the edible, non-bitter, and highly nutritious seeds, as the fruit flesh was bitter and became papery at maturity. Under domestication, selection for larger seeds probably inadvertently resulted in selection for larger fruits. Harvesting of immature fruits for food resulted in encounters with mutants having non-bitter flesh. Cutting of mature fruits for their seeds resulted in encounters with and preservation of seeds from mutants having non-bitter, starchy flesh. Presumably, selection for non-lignified rinds and improved quality of the mature fruits followed. The English-language terms for C. pepo--pumpkin, squash, gourd, and marrow-have often been used indiscriminately. Sturtevant (Hedrick 1919) adequately reviewed the literature concerning the usage of the terms pumpkin, squash, and gourd. Some repetition would be valuable as these terms will be used to some extent here in distinguishing the various forms. The term squash is derived from a North American Indian word, asq, plural asquash, whose root meaning is something eaten when immature or raw (Trumbull 1876). On the other hand, the word pumpkin is derived from the Latin pepo and Greek pepon, which implies roundness and larger size (Hedrick 1919). Thus, the modern common usage of the term summer squash for the immature fruits of C. pepo and the term pumpkin for the mature fruits of large, round-fruited forms of Cucurbita admirably fits the ancient usages of these terms. The modern term winter squash is applied to palatable mature fruits that are not round or nearly so. Various suggestions in the scientific literature for distinguishing a squash from a pumpkin by species (Castetter and Erwin 1927) or by eating quality and method of culinary preparation (Whitaker and Bohn 1950) deviate from the original meanings and have not been incorporated into vernacular usage. The term gourd will be used here in the same sense as is used in the U.S.A.: unpalatable forms of C. pepo and other cucurbit species. The word marrow, a shortened version of "vegetable marrow," is used in the U.K. to denote the immature fruits of C. pepo. The term vegetable marrow is reserved for a particular group of summer squash, a group that was popular in the U.K. (see below). As far as has been determined, all of the various pumpkins, squashes, and gourds of Cucurbita pepo intercross freely. And yet, as pointed out by Sturtevant (Hedrick 1919) and Bailey (1929), some cultivars and groups of cultivars have retained their distinguishing characteristics for centuries. The objective of my review is to trace the history, possible origins, modern development, and present status of the culinary groups of C. pepo. HISTORICAL RECORDS OF THE CUCURBITA PEPO CULINARY GROUPS The earliest written reports we have of Cucurbita are accounts of the first explorers of the Americas. These early records, reviewed by Sturtevant (Hedrick 1919) and by Coulter (1943), do not appear to have contained detailed descriptions of fruit types. Some 50 yr after the voyage of Columbus, descriptions and illustrations of Cucurbita began to appear in botanical herbals of Europe. They include the first accurate detailed descriptions in existence of the various forms of C. pepo. These descriptions were written some 100-200 yr before the Linnean system of botanical classification was proposed, and many different Latin names were ascribed to the

426 ECONOMIC BOTANY [VOL. 43 same or similar forms over the years. Lagenaria, a genus containing gourds of very many shapes, was included by many early botanists, including Linnaeus, in Cucurbita (Bailey 1929). Thus, the accounts of the various forms classified as Cucurbita must be interpreted with particular caution. Fortunately, flower color was an important characteristic to these botanists, and in many accounts the mention of white (for Lagenaria) or yellow (for Cucurbita) flower color allows classification with certainty as to genus. Some vernacular accounts have also been a potential source of confusion. For example, pumpkins were variously referred to as pompions, pompeons, melons, millions, and yet other terms. Even the same form or cultivar often had more than one vernacular name, a condition existing to this day. In addition, the same name was often applied to completely different cultivars. An important example of the latter case was the use of the term crookneck for a form of C. pepo summer squash and for a form of C. moschata (Duch. ex Lam.) Duch. ex Poir. winter squash. Fortunately, most of the early botanical drawings were so detailed that the particular species of Cucurbita to which the depicted form belonged is easily determined, based on the distinguishing characteristics of the cultivated Cucurbita species enumerated by Whitaker and Bohn (1950). To these drawings can be added some realistic paintings from the 16th and 17th centuries that included Cucurbita fruits (Zeven and Brandenburg 1986) as well as detailed literary accounts, notably the Historia plantarum universalis (Bauhin 1651), which was completed by 1613 (Morton 1981). These drawings, paintings, and literary accounts also enable us to trace, to a greater or lesser extent, the history of each of the C. pepo groups, as follows: Pumpldn The pumpkins, probably the oldest and most diverse group of C. pepo, have been considered to be the typical group of the species (Bailey 1929; Greben~ikov 1958; Paris 1986b). They were the first C. pepo to be depicted in the botanical herbals (Bock 1546; Fuchs 1542) and were subsequently represented in great variety, including forms having fruits that were spherical, oblate, oval, obovate, longitudinally grooved and non-grooved, and large and relatively small (Bauhin 1651; Dalechamps 1653; Dodoens 1554, 1616; Gerard 1597; Gerarde and Johnson 1636; Lobelius 1576; Mattioli 1558, 1586; Parkinson 1640; Tabernaemontani 1664). An illustration by Fuchs (1542) may represent 'Small Sugar', a cultivar that is still very much in commerce. In realistic paintings of vegetable markets in the Netherlands and Flanders by various artists of the 16th and 17th centuries, as many as eight different forms of C. pepo pumpkins were depicted (Zeven and Brandenburg 1986). Pumpkins also became denizens of English gardens (Miller 1732; Parkinson 1640; Plat 1660; Quintinye 1710), and forms subsequently found in North America were compared with those already being grown in the U.K., both favorably (Beverley 1705; Josselyn 1672) and unfavorably (Strachey 1612). Numerous pumpkins were listed and described in accounts of then contemporary cultivars in the U.K. (Corbett 1833; Mawe and Abercrombie 1797) and North America (Burr 1863; Castetter and Erwin 1927; Fessenden 1837; Goff 1888; Gregory 1893; Tapley et al. 1937). Zhiteneva (1930) presented photographs of

1989] PARIS: CUCURBITA PEPO 427 fruits from land races of C. pepo from Turkey and Mexico, including many pumpkins. Pumpkins are consumed when the fruits are fully mature. Their fruit rinds are not lignified, enabling easy slicing of the fruits. This characteristic apparently was important to Native Americans, for it has been suggested (Kay et al. 1980; Whitaker and Cutler 1965) that it was common practice to slice the fruits into strips and then hang the strips to dry, for preservation of the flesh. Scallop The scallops are one of the oldest cultivar groups of C. pepo. A drawing of a scallop form, perhaps representing the still extant cultivar 'White Bush Scallop', appeared in the herbal of Dodoens (1554). Bauhin (1651) reported seeing "Cucurbita clypeiformis sive Siciliana, Melopepon latus ~ nonnullis vocata" in 1561 and again in 1606. The accompanying illustration appeared in other herbals, including that of Gerard (1597), who labelled it "Great Buckler Pompion.'" This form is perhaps the still extant cultivar known nowadays as 'Golden Bush Scallop' or 'Golden Custard'. In describing the Great Buckler Pompion, Gerard stated that it was new in England even though the other forms depicted, all round- or nearly round-fruited, were common. Thus, the scallops may have been disseminated in Europe a few years later than the pumpkins. In contrast to the pumpkins, the scallops are grown for culinary use of their immature fruits. The consumption of immature fruits of C. pepo is an ancient practice (Cutler and Whitaker 1961); such use by Indians was described in 17th century accounts of North America (Williams 1643; Wood 1634). The different culinary usage of pumpkins and scallops by Indians was described by Beverley (1705). The scallops are woody and inedible as mature fruits, but are firm and quite pleasant as young fruits. From literary accounts in the U.K. (Corbett 1833; Mawe and Abercrombie 1797) and U.S.A. (Burr 1863; Goff 1888; Neill 1851; Tapley et al. 1937) it can be concluded that the scallops were grown continually over a long period of time. Alefeld (1866) assigned the Latin name C. pepo clypeata to the scallops. Acorn Whitaker (1947) reproduced an illustration of an acorn form appearing in the Tabernaemontanus edition of 1591. This and other illustrations of acorns also appeared in Gerard (1597), Bauhin (1651), and Tabernaemontani (1664). The acorn labelled by Gerard "Cornered Indian Pompion" and by Bauhin and Tabernaemontani "Pepo indicus minor angulosus" appears to have had lower shoulders and shallower furrows than those of modern acorn cultivars. A fruit of fuller acorn shape, including squarer shoulders and deeper furrows was depicted and labelled by Bauhin, who observed it in a garden in 1607, as "Cucurbita clypeata affinis ~ pediculo in conum desinens,'" and by Tabernaemontani (1664) as "Pepo minor Indicus IV" (the illustration may have first appeared in 1591 edition). An interesting third acorn form depicted was the "Mushrome wilde Gourde" of Gerard or "Cucurbita capitata" of Bauhin and Tabernaemontani; the illustration is suggestive of the 'Sweet Dumpling' cultivar. Although the fruits are 10-ribbed and pointed at the apical end like those of other acorns, this cultivar is unusual in having a depressed, concave peduncle end and dark green stripes in the furrows of the fruits. An acorn fruit having a blunt blossom end and rather rounded

428 ECONOMIC BOTANY [VOL. 43 shoulders, but strongly furrowed, was illustrated by Rumpf(1757). Dickson (1807) mentioned cultivars having conical and turbinate "torulose" fruits. Both Metzger (1841) and Roemer (1846) described several forms having fruits that were 10- ribbed, slightly longer than wide, and convex at their apical end. Naudin (1860) depicted in color an acorn fruit shaped much like those of modern cultivars: it was distinctly furrowed, shouldered, conical, and pointed with a small blossom scar. Perhaps the earliest mention of a squash cultivar by the name of Acorn in North America was in the listing of squash forms by Fessenden (1837). However, in the mid- 19th century, Acorn was synonymous with Turban or Turk's Cap (Burr 1863), a cultivar of C. maxima Duch. ex Lam. According to Castetter and Erwin (1927), the first well-known acorn cultivar grown in North America was a dark green-fruited form by the name of 'Table Queen', which was listed by a seed company for the first time in 1913. Prior to its introduction, this cultivar or its forerunner was reportedly grown by several farmers near Des Moines, IA, under the name Danish squash. A similar but inferior form was said to have been raised by the Arikara Indians of North Dakota. Crookneck The crooknecks do not appear to have been described by European herbalists. Sturtevant (Hedrick 1919) wrote that the crookneck may have been observed by the French explorer Champlain in 1605. Although he did not cite his source of information, Sturtevant did carefully distinguish between the winter crookneck (C. moschata) and the crookneck of C. pepo. In the British gardening compendia of Miller (1768), Hanbury (1771), and Mawe and Abercrombie (1797), warted forms were described as being viney and highly branched, as having fruits that were oblong, round, flat, or bottle-shaped, and as being used in America for culinary purposes when the fruit was immature. Duchesne (Lamarck 1786) mentioned elongate, warted, yellow gourds ("les barbarines jaunes"). Likewise, Willdenow (1797) described warted, yellow, variable forms of elliptical, club, or elongate shape. The peculiar crookneck characteristic was not mentioned by name by any of these authors. However, the crooknecks and the straightnecks (see below) are the only groups of cultivars (excluding some ornamental gourds) that, at least nowadays, are comprised mainly of forms having fruits that are both warred and yellow. It may also be important to note that the botanist and plant explorer T. Nuttall (1818) mentioned finding a "warted squash"... "'Cultivated also by the Indians of the Missouri to its sources" and that 10 yr later a crookneck of C. pepo first appeared in a seed catalog. The earliest use of the word crookneck for a form of C. pepo appears to be that referred to by Sturtevant (Hedrick 1919) and Tapley et al. (1937), who stated that a cultivar by the name of'summer Crookneck' first appeared in a North American seed catalog as early as 1828. Other early North American listings were those of Fessenden (1837), Bridgeman (1840), Wood (1849), Burr (1863), and White (1868). The first references to a crookneck form by name in Europe may be the listings of Corbett (1833) and Seringe (1847). The latter listed "Courge crochue" as a little-known form eaten only when young. Naudin (1856) described two crookneck forms, both warted, the bush-habit "Courge Coutors" or "Crook-neck des Amrricains" and the vine-habit "Courge Polk," the latter of which was illustrated. Alefeld (1866) assigned the Latin name C. pepo

1989] PARIS: CUCURBITA PEPO 429 torticollis to the crooknecks. All modern crooknecks are of bush habit, but according to Tapley et al. (1937), viney forms of crookneck were once common. Straightneck Tournefort (1700) depicted a warted, oblong, slightly necked form in his herbal. In the British gardening compendia (Hanbury 1771; Mawe and Abercrombie 1797; Miller 1768), bottle-shaped warted forms used for culinary purposes in North America are mentioned; Duchesne (Lamarck 1786) mentioned elongate, warted, yellow "barbarines"; and Willdenow (1797) described warred, yellow clubshaped squash. However, it is most peculiar that, although the crooknecks appeared by 1828, the straightnecks were not mentioned by early and mid-19th century writers and are absent from the review of American garden plants by Burr (1863) and in the reviews of Cucurbita cultivars by Goff (1888) and Gregory (1893). The lone cultivar of the straightneek type described by Tapley et al. (1937) is 'Giant Summer Straightneck', reportedly a selection from 'Giant Crookneck' and introduced in 1896. In addition, the well-known cultivar 'Early Prolific Straightneck', introduced in 1938 by Ferry-Morse, was reportedly a selection from 'Summer Crookneck' (Minges 1972). Vegetable marrow Both Bailey (1929) and Whitaker (1947) cited an illustration from the herbals-- the "Pepo maximus oblongus" in Gerarde and Johnson (1636) and a form of the same name in Tabernaemontani (1664), respectively--of what each author considered to be a vegetable marrow form. However, the fruits depicted were not noticeably broader toward the blossom end, a characteristic diagnostic of the vegetable marrow group (Paris 1986b; Fig. 6, 14). Sturtevant (Hedrick 1919) suggested that these forms may have been closely allied to the 'Long Yellow Pumpkin'. Another possibility is that these were the forerunners of the large pumpkin cultivars 'Sandwich Island' and 'Tours' (described by Robinson 1885 and Tapley et al. 1937). A possible forerunner of the vegetable marrow group was a bush form having fruits that were hardly longer than wide but were noticeably tapered, broader at the blossom end. This form was illustrated and labelled by Gerard (1597) "Great Wilde Pompion," by Tabernaemontani (1664) "Cucurbita longa," and by Bauhin (1651) "Cucurbita indica minor." Several other possible candidates as forerunners of the vegetable marrow group also appear in these works. In spite of their apparently long history, the vegetable marrows were not described or even mentioned in the late 18th and early 19th century British gardening compendia (Hanbury 1771; MacPhail 1819; Mawe and Abercrombie 1797; Miller 1768) or in a botanical account of Cucurbita from the same era (Willdenow 1797). Sanders (1910) recounted that little was known of the early history of the vegetable marrow. He mentioned a paper published in Transactions of the Royal Horticultural Society in 1816 by J. Sabine, who reportedly suggested that the vegetable marrow was brought to England from Persia, but this claim has not been substantiated and seems unlikely because several incipient forms existed 200 yr earlier in Europe. Tapley et al. (1937) reported that the vegetable marrow first appeared in a North American seed catalog in 1824. Corbett (1833),

430 ECONOMIC BOTANY [VOL. 43 writing in England, mentioned a "long bush" form of squash. The vegetable marrow was listed as a little-known form by Seringe (1847). Naudin (1856) described and translated it as Courge ~t la Moelle. It was also mentioned in North America by Fessenden (1837), Bridgeman (1840), Burr (1863), and White (1868). Both bush and vine forms of vegetable marrows existed (Tapley et al. 1937). Much of the great diversity of vegetable marrow cultivars is probably attributable to the development of this form in the U.K. during the 19th century. Cocozelle According to Webster's third new international dictionary (Gove 1976), the word cocozelle is probably derived from an Italian dialect word, cocuzza, for squash. The word cocuzza was used for the immature fruits of an extremely longfruited form of Lagenaria, which is still grown today for use as a summer squash. A long-fruited form of Cucurbita pepo, described as 26 units long and no greater than 8-10 units broad, was observed growing in a garden in the early 17th century by Bauhin (1651), his "Zucha major longa." This may have been a forerunner of the cocozelle. The cocozelle, as "Courge Coucouzelle," was first mentioned by Seringe (1847). Excellent descriptions of the cocozelle from both Europe and North America were provided by Naudin (1856) and Burr (1863), respectively. Fine illustrations with slightly different spellings for 'Cocozelle' accompanied the writings of Robinson (1885) and Tamaro (1916). The 'Cocozelle' cultivar appears to be an important example of forms that were generally referred to in the U.S.A. as Italian vegetable marrows (Tapley et al. 1937). Another cocozelle form is the "Courge blanche trrs allongre" of Naudin (1856), which is probably the same as the 'Long White Bush' illustrated by Robinson (1885) and described by Goff (1888) as 20" long and 4" in broadest diameter. The fruits of PI 177370 (from Turkey) and PI 379307 (from Yugoslavia) adhere closely to the descriptions and illustration of this form. PI 261610 (from Spain) is a dark-fruited cocozelle form. Zhiteneva (1930) presented photographs including several cocozelle fruits from Asia Minor. Zucchini The word zucchini is the Italian plural for summer squash. The "Zucha major longa" of Bauhin (1651) may also have been a forerunner of the zucchini. The zucchini was first mentioned by Naudin (1856) as zouquette or zouquetti from the port of Marseilles, France. The first full description of a zucchini was also by Naudin (1856), his "Courge de Larnaca." This form was described as having fruits almost uniformly cylindrical in shape, 40-50 cm long, and 10-12 cm wide. The fruits were reportedly smooth and yellowish white and the plants viney, semibush, or bush. A later description of dark-colored, bush and vine zucchini cultivars appeared in Italy (Tamaro 1916). The zucchini was also grown in Asia Minor, as evidenced by the photographs of Zhiteneva (1930). According to Tapley et al. (1937), the zucchini was introduced into the U.S.A. from Italy during the 1920s and within a decade it constituted a distinct group. Shifriss (1981) described the origin of the well-known cultivar 'Fordhook Zucchini' as Naples, Italy, followed by inbreeding and selection in the U.S.A. 'Fordhook Zucchini' was introduced by the Burpee Seed Co. in 1942 (Minges 1972).

1989] PARIS: CUCURBITA PEPO 431 Other culinary forms of Cucurbita pepo Other fruit shapes exist in C. pepo that cannot readily be included in any one of the horticultural categories above. Some of these are intermediate forms. For example, many so-called zucchini hybrids have long, noticeably tapered, nonbulbous fruits. Such hybrids are in reality F~ s between vegetable marrows and zucchinis; thus their intermediate appearance. Other cultivars, such as 'Delicata', are more or less unique and are not readily classifiable in any one modern group. 'Delicata' was first introduced by a seedsman in the U.S.A. in 1894 (Tapley et al. 1937), but a fruit very much like those of this cultivar was illustrated by Naudin (1856). ORIGINS OF THE CULTIVAR GROUI~ Recent reports on fruit rind development (Schaffer 1983), allozyme variation (Andres 1987; Decker 1985; Ignart and Weeden 1984), and seed morphology (Decker and Wilson 1986) provide useful information concerning evolution and systematics of the groups of C. pepo. Some hypotheses have been developed regarding genetic relationships within the species (Decker 1985; Decker and Wilson 1986). As fruit shape is under multigenic control (Sinnott 1935), it too has some value in considering these relationships. The present consideration of the origin of the groups is limited to accounting for the majority of their extant cultivars. Any or all of the groups might contain cultivars that were derived at different times and/or at more than one location and/or by more than one means. Alternative pathways of derivation for each of the groups are therefore not mutually exclusive. What presently appears to me to be the more likely alternative for the majority of the extant cultivars is summarized pictorially in Figure 17. Pumpkins and scallops As the pumpkins (Bock 1546; Fuchs 1542) and shortly thereafter the scallops (Dodoens 1554) were the first to be depicted in the botanical herbals, it can be concluded that both groups were brought to Europe from America. The pumpkins must have existed in a variety of forms at the time of the Discovery, as attested to by the records of J. Bauhin (1651) and paintings of 16th century artists (Zeven and Brandenburg 1986). At least two scallop cultivars were introduced into Europe by 1561 (Bauhin 1651; Dodoens 1554). Recently, a hypothesis was developed from studies ofallozyme variation (Decker 1985; Ignart and Weeden 1984) and seed morphology (Decker and Wilson 1986) that there exists a fundamental division within C. pepo, with the pumpkins and scallops lying on opposite sides of that division. The great differences in fruit shape and culinary adaptation of these two groups would also suggest that they are genetically and evolutionarily distant from one another. Some rare pumpkin cultivars have lignified rinds and even warts. Expression of lignified rind is conditioned by the dominant gene Hr (hard rind) and of warts by the dominant gene Wt. Both Hr and Wt must be present for wartiness to develop (Schaffer et al. 1986a). Primitive forms from Mexico, exemplified by PIs 442315 and 442325, have smooth but thick lignified rinds; these forms may be representative of an earlier stage in evolution under domestication, an early off-

432 ECONOMIC BOTANY [VOL. 43 Fig. 9-16. Representative mature (40-5O d past anthesis) fruits of the eight Cucurbita pepo cultivar groups grown for culinary purposes. Fig. 9, Pumpkin 'Spookie'. Fig, 10. Crookneck 'Early Yellow Crookneck'. Fig. 11. Straightneck 'Early Prolific Straightneck'. Fig. 12. Scallop 'Benning's Green Tint'. Fig. 13. Acorn 'Mammoth Table Queen'. Fig. 14. Vegetable Marrow 'Sihi Lavan'. Fig, 15. Cocozelle 'Coeozelle'. Fig. 16. Zucchini 'Black Zucchini'. Not to scale.

1989] PARIS: CUCURBITA PEPO 433 shoot, or a separate line of domestication from the pumpkin line. The South African cultivar 'Little Gem' is a miniature pumpkin having a smooth, thin, but lignified rind. 'Nantucket', a pumpkin cultivar of the 19th century (Burr 1863; Goff 1888), had warts. The reportedly very high quality of 'Little Gem' and 'Nantucket' indicates that these pumpkins are well advanced from primitive stocks. Their hard rinds would suggest that they were derived through hybridization between pumpkins and hard-rinded forms. Acorn The acorn appeared in botanical literature beginning with the Tabernaemontanus edition of 1591 (Whitaker 1947). This is some 30-50 yr after the first records of the pumpkins and scallops in Europe. The timing here is perhaps significant: the acorns may have been derived through chance crossing between the pumpkin and scallop groups (Fig. 17). A cross between a pumpkin and a scallop could account for the acorns having the non-lignified rinds and the satisfactory mature fruit eating quality of the pumpkins and the "cornered" shape of the scallops. Indeed, the similarity of the shape between the acorns and scallops is attested to in an unusual way in the herbals: in Gerard's 1597 herbal the "Cornered Indian Pompion" is illustrated as an acorn form but in a later edition (Gerarde and Johnson 1636), the "Cornered Indian Pompion" is illustrated as the scallop form of Dodoens (1554). Dickson (1807) listed "conical-" and "turbinate-" fruited forms among the scallop cultivars. Consequently, the possibility exists that acorns may have been observed by others through the centuries but were included with the scallops. The acorn depicted by Naudin (1860) had an unusual color pattern: cream with narrow green stripes in the furrows, as in 'Sweet Dumpling' and 'Delicata'. The furrows (and dark stripes) correspond to the position of the l0 main carpellary vascular bundles (Bailey 1937). Naudin (1856) considered 'Delicata' and other unusual fruit types he obtained to have originated from chance crossing between plants of different groups, specifically between a scallop (or acorn?) and a warted gourd. Another probable derivative of chance crossing between forms of different groups in Naudin's research plots is the 'Cocoanut' cultivar of Gregory (1893), which was introduced into the U.S.A. in 1869 (Tapley et al. 1937). This unusual, furrowed, striped, and bicolor cultivar was probably derived from crossing of Naudin's striped acorn form and a bicolor gourd (Paris 1986a). In a recent study of seven polymorphic allozyme loci (Decker 1985), most of the alleles predominant in 'Table Queen' were also predominant in the scallops; however, at two loci, the predominant alleles in 'Table Queen' differed from those of the scallops and were the same as those in two pumpkin cultivars. The hypothesis for the derivation of the acorns from the pumpkins and scallops seems consistent with the botanical record and the present knowledge of genetic relationships. However, further investigation is warranted. Of particular interest would be attempting to resynthesize acorns from crosses of modern pumpkins and scallops. Crookneck and straightneck The accounts of Sturtevant (Hedrick 1919) and Tapley et al. (1937) suggest that crooknecks existed long before 1828. This and other literature (Nuttall 1818)

434 ECONOMIC BOTANY [VOL. 43 Wild Gourd 8 Ornamental Gourd Crookneck Straightneck Scallop Cocozelle Acorn Pumpkin Fig. 17. A proposed scenario for the origin of the modem, extant forms of the cultivar groups of Cucurbita pepo grown for culinary purposes. would also suggest that the appearance of the crooknecks in recorded history was delayed because they were cultivated in interior regions of North America, which were inaccessible to early explorers. Thus, the possibility exists that this group is an ancient and separate line of domestication, like the pumpkins and the scallops (Fig. 17). Unlike the other two groups, the crookneck apparently was not widely distributed nor did it consist of a number of distinct variants. As crooknecks have been cultivated continuously in North America at least since 1828, it seems safe to assume that the modem crookneck cultivars are the direct descendants of those cultivated over 150 yr ago. The origin of the straightnecks of today appears to be quite recent, American, and separate from the incipient form depicted by Tournefort (1700). To the best

1989] PARIS: CUCURBITA PEPO 435 of my knowledge, the straightnecks have been and are grown almost exclusively in North America. Both the crooknecks and straightnecks are characterized by warted rinds (Hr, Wt) and yellow immature fruit color (Y), characteristics occurring in relatively few other culinary forms ofc. pepo. Also, the modern straightnecks generally resemble the modern crooknecks with regard to foliage and plant characteristics. Representative cultivars of the two groups were found to differ in the predominant allele for two of seven allozyme loci (Decker 1985), but they clustered in canonical variate and in principal component analysis of seed morphology (Decker and Wilson 1986). There seems to be no other reasonable conclusion than that the extant straightnecks are relatively recent derivatives of the crooknecks (Fig. 17). Vegetable marrow, cocozelle, and zucchini Derivation of the vegetable marrows from the pumpkins was suggested by Decker (1985) from allozyme analysis data; this suggestion is c~nsistent with the herbal record. Conceivably, the vegetable marrows, cocozelles, and zucchinis could owe their origin to the hard-rinded primitive cultivars from Mexico. PI 442294 from Mexico has fruits about 30 cm long and 9 cm wide, resembling in dimensions Bauhin's "Cucurbita major longa." The fruit is very much like other zucchinis; it is cylindrical, only slightly tapered, with a light peduncle (d/d) and dark (L-l~-- L-2/--) (Paris and Nerson 1986) green fruits. Allele L-1 is relatively infrequent in C. pepo, occurring mainly in the zucchini group. Most pumpkins carry gene D for dark stems and peduncles, but the Mexican forms from the same collection that I have grown are all d/d. The plants of PI 442294 are typical of those of others in the collection: viney, late, rank, coarse growth, and strongly male--in short, very primitive. Thus it seems likely that PI 442294 is not an import, but that it is truly Mexican. It seems possible that the vegetable marrows, cocozelles, and zucchinis could have been derived from primitive Mexican pumpkins through accumulation of genes for long fruit shape, and then brought to Europe by early explorers, improved in European gardens, and then carried to America in the 19th century. However, from this hypothesis it would not be easily asserted that the vegetable marrows were derived from more advanced, typical pumpkins, as all vegetable marrows have lignified rinds whereas the pumpkins have non-lignified rinds. Another possible scenario for the derivation of the vegetable marrows, cocozelles, and zucchinis of.today is through chance crosses between non-lignified pumpkins and a somewhat elongate ornamental gourd, such as 'Pear', in Europe (Fig. 17). 'Pear' was illustrated by Lobelius (1576). This illustration reappeared in Gerard (1597), who labelled it "Peare Fashioned Coloquintida," in Dodoens (1616), who labelled it "Cucurbita silvestris," and in Tabernaemontani (1664), who labelled it "Coloquint das Beiblein." The length to broadest width ratio of 'Pear', approximately 1.5, is high relative to other C. pepo illustrated prior to 1600. And as in the vegetable marrows and cocozelles, the fruits of 'Pear' are distinctly broader at the apical (blossom) end and narrower at the basal (peduncle) end. Chance crossing of a pumpkin and 'Pear', followed by introgression of gene Hr and some genes for elongate fruits from 'Pear' into the pumpkin, could have occurred in Europe well prior to 1600, and through selection for longer and more palatable fruits the vegetable marrows, cocozelles, and zucchinis were derived.

436 ECONOMIC BOTANY [VOL. 43 'Pear' has broad, dark stripes alternating with narrow, light stripes. As opposed to the striping of some acorns and their allies, in 'Pear' the light stripes correspond to the main carpellary vascular tracts; this type of striping, which also occurs in other ornamental and wild C. pepo gourds, is rare in culinary forms. However, this pattern occurs in the fruits of some cocozelle (Fig. 7, 15) and vegetable marrow cultivars. Shifriss (1965) presented a photograph of bicolored, long-oblong fruits he obtained from Mexico. The bicolor characteristic is conditioned by rare mutants of gene B + and occurs in the ornamental gourd cultivar 'Bicolor Pear'. In attempting to explain the source of the bicolor characteristic in the Mexican material, Shifriss suggested two alternative possibilities: that the mutation occurred in this peculiar Mexican cultivar or that this cultivar is a derivative of hybridization between a bicolor gourd such as 'Bicolor Pear' and edible cultivars. 'Gray Zucchini' (in spite of its name, a cultivar of the vegetable marrow group) and 'Black Zucchini' are similar to pumpkins with respect to allozyme allele frequencies (Decker 1985). However, for one allozyme (Idh-lg) examined, the predominant alleles are not the same; for this locus, the predominant allele in 'Gray Zucchini' and 'Black Zucchini' matches that of'pear'. Of the six allozyme loci examined by Ignart and Weeden (1984), the vegetable marrow and zucchini were found to have different predominant alleles from the pumpkins in two to four, depending on the cultivar. As a whole, present knowledge seems to be more consistent with the hypothesis that the vegetable marrows, cocozelles, and zucchinis were derived through hybridization rather than directly from the pumpkins. The feasibility of the hybridization hypothesis can be tested by crossing a pumpkin such as 'Small Sugar' with 'Pear' and attempting to isolate long-fruited lines in filial or backcross generations. MODERN DEVELOPMENT AND PRESENT STATUS OF THE CULTIVAR GROUPS Until approximately 60 yr ago, Cucurbita cultivars were characterized by much variability, attributable to the outcrossing tendency of this species. With increased demand for uniformity and refinement of various characteristics--e.g., earliness, fruit size, color, shape, and quality--highly homozygous, true-breeding cultivars and strains were developed (Bushnell 1922; Haber 1928; Scott 1934; Sinnott and Durham 1922). A number of these uniform, homozygous strains became important commercial cultivars. During the past 35 yr many inbreds have been used as parents in the development of F1 hybrids. These hybrids, although heterozygous, are uniform and homogeneous. The modern evolution of edible C. pepo under domestication has been following two divergent courses: increased adaptation to production and use as summer squash and increased adaptation to production and use as pumpkins and winter squash. These two courses have been divergent because of the different demands and preferences that growers and consumers have for the immature fruits as opposed to the mature fruits of C. pepo. The forms grown as summer squash have fruits that diverge strongly from a 1:1 ratio of length to broadest width. These are the various elongate-fruited forms--the crooknecks, straightnecks, vegetable marrows, cocozelles, and zucchinis--and the flattened forms--the scallops. The forms grown for their mature fruits, the pumpkins and acorn squashes, have a

1989] PARIS: CUCURBITA PEPO 437 length to (broadest) width ratio of very nearly 1:1. The association between length to width ratio and the stage at which the fruits are consumed is generally true in other economically important Cucurbitaceae. The deviation from the 1:1 ratio can be attributed to the consumer desire for a narrow diameter seed cavity in immature fruits. Divergent evolution under domestication for fruit shape as a function of culinary use is still occurring in C. pepo today, witnessed by the increasing importance and active breeding of summer squash of the zucchini type in many regions. Likewise, summer squash differ in plant architecture from pumpkins and winter squash, as described below. Summer squash As summer squash need to be picked several times weekly in order for their fruits to be of high quality and for subsequent fruits to be produced, much labor is required for harvesting, and this labor is often expensive and at times of limited availability. Summer squash fruits are tender (a consumer demand) and thus mechanical harvesting has not been possible. Therefore, a strong premium exists on cultivars which can be harvested quickly. Vining forms of summer squash, once common, are now rare, because bush habit facilitates harvesting. Not surprisingly, long and fiat fruit shapes are associated with bush habit in C. pepo. The past 35 yr have been witness to further improvements in plant architecture and fruit characteristics of summer squash. Open growth habit--a characteristic involving lack of branch production and low (more horizontal) petiole angle-- results in saving much of the time required for harvest (Baggett 1972). Many modern cultivars carry this important improvement to at least some degree. Another characteristic of importance in summer squash is smoothness of the foliage. The foliage of C. pepo is prickly. At harvest, this results in scratching of the fruit, which detracts from appearance and marketability. During the past 25 yr some efforts have been directed to developing cultivars having smoother foliage. Color is an important component of appearance and, as with other vegetables and fruits, intense coloration is a positive attribute (Higgins 1986; Rodriguez and Sanchez 1983). Two complementary dominant genes, designated L-1 and L-2, are required for expression of intense fruit color in summer squash (Paris and Nerson 1986). Of the summer squash groups, the zucchini is the most advanced by far in incorporating these horticulturally valuable characteristics. Of particular recent interest and potential usefulness in the breeding of summer squash has been the precocious yellow fruit characteristic, which is conditioned by a single gene, designated B (Shifriss 1965). In concert with L-1 and L-2, B imparts an attractive, intensely yellow color to the squash. The intensely yellow fruits can be harvested approximately 25% faster than green fruits (Paris et al. 1986). Furthermore, the fruits tend to be slimmer (Shifriss 1966) and to grow more slowly (Paris et al. 1983, 1986; Schaffer and Boyer 1984), which are distinct advantages for marketing in Europe and the Middle East, as in these regions there exists a strong preference for small fruits. Breeding for disease resistance in C. pepo has focused on the summer squash and the transfer of genes for resistance from C. moschata and wild Cucurbita (Munger and Washek 1983; Whitaker and Robinson 1986); as in most cases sources of resistance within C. pepo are unknown or non-existent (Whitaker and

438 ECONOMIC BOTANY [VOL. 43 Robinson 1986). The search for sources of resistance and the difficulties encountered in interspecific crossing make such breeding time consuming and expensive. Such investment is likely to be made initially only for the forms of highest economic value. As the zucchini is of economic importance worldwide, and the crookneck and straightneck of importance in the U.S.A., where capable personnel and sufficient capital for investment exist, these are the likely groups into which the genes from other species will be incorporated. The infiltration of genes from other species of Cucurbita into C. pepo might further contribute to the divergence within the summer squash and between the summer squash and the pumpkins and acorns. The scallop cultivars are grown today mainly by home gardeners and have some minor commercial importance in the U.S.A. Until recently all cultivars were quite primitive with respect to several horticulturally important characteristics: the plants were spiny, of closed growth habit, and late bearing. Some breeding has been conducted in the U.S.A., and this has resulted in the release of hybrid cultivars that are earlier and more productive and have a more open growth habit, less spiny foliage, and more intensely colored fruits than those of their predecessors. In the Middle East and North Africa, a common method of culinary preparation of summer squash is to stuffit with meat and rice. As the scallops by their broad shape are adapted to being stuffed, there may be an opening for expansion of scallop cultivars, at the expense of the vegetable marrows, in this region. The crooknecks and straightnecks have retained their popularity in the U.S.A. Nearly all modern cultivars are hybrids, bred in the U.S.A. for use in the U.S.A. Modern hybrids contain important improvements such as small plant size and open growth habit for easy harvesting, clearer yellow fruit color, less wartiness, stronger female tendency, and higher yields. The vegetable marrow cultivars appear to have undergone little improvement during the last century. Most modern vegetable marrows are bush forms; as a group they are rather primitive horticulturally, most cultivars having very spiny foliage and a very closed growth habit. The cultivars 'Caserta' and 'Gray Zucchini' stand apart from other vegetable marrows by their open growth habit and fruits that are at the extreme upper limit for length:width ratio of the vegetable marrows, 3.0 (Paris 1986b). The vegetable marrow apparently never achieved popularity in North America, but in the Middle East and North Africa it is the most common form of summer squash. Recently, some American companies have begun breeding for the Middle Eastern market and released vegetable marrow hybrids that are generally improved for foliage characterstics. For years these companies have utilized inbreds of 'Caserta' in crosses with zucchinis to produce "zucchini" hybrids. The cocozelle is well represented in countries bordering the Mediterranean Sea, e.g., 'Cocozelle' (originally from Italy), PI 177370 (Turkey), PI 261610 (Spain), and PI 379307 (Yugoslavia). A very long, striped cocozelle is also the predominant cultivar in Hungary (H. Nerson, pers. comm.). The cocozelles underwent improvement and development, including production of hybrid cultivars, in the U.S.A. during the middle part of this century (Minges 1972). Two new, highly advanced, hybrid cocozelles were recently released by the Hung Nong Seed Company of South Korea.

1989] PARIS: CUCURBITA PEPO 439 The zucchini is today the most advanced group of summer squash in horticulturally important foliar characteristics and is the most economically important and extensively grown C. pepo cultivar group in the developed countries. Most zucchini cultivars are hybrids. Much of the improvement of this group has been by American breeders through inbreeding and selection within variable forms from Italy during the last half century. During the last 25 yr there has been a strong trend toward breeding more productive, pure zucchini hybrids having very intensely colored fruits and relatively small, single stem, open habit plants with smooth foliage. There is also a general trend for replacement of the vegetable marrow, cocozelle, and less advanced zucchini cultivars with these modern, intensely green zucchinis. A similar trend for replacement of the crookneck and straightneck cultivars by the modern intensely yellow zucchinis may also have begun. In countries where summer squash is relatively new as a large-scale commercial crop, the type grown is almost always the zucchini. Worldwide, the zucchini will probably continue to be the most economically important group of C. pepo. Pumpkins and winter squash The demands made on the forms used for their mature fruits, the pumpkins and acorns, contrast with most of those made on summer squash. Bush habit does not carry a high premium, except for home garden cultivars and in the development of semi-bush F1 hybrids. These hybrids, obtained by crossing a vine form with a bush form, have high fruit quality (like the vine cultivars) and bushy growth, which facilitates cultural and harvesting operations. Characteristics of importance in mature C. pepo fruits are puncture resistant, but non-lignified rind, intensely colored rind and flesh, and thick, fine-grained, high-quality flesh high in dry matter and carotene. The complementary action of genes B and L-2 impart orange flesh color and can increase carotene content by as much as 15-fold (Paris et al. 1985; Schaffer et al. 1986b). Pumpkins of C. pepo remain popular today in many parts of the world for consumption of the flesh in a huge variety of ways (Jabs 1979; Ralston and Jordan 1977), seed consumption, ornament, and Halloween decoration. Acorn squash is common in American supermarkets, is considered to be of good quality, and is also prepared in a wide variety of ways (Ralston and Jordan 1977). Further improvements in fruit characteristics, such as thicker and more nutritious flesh, can be expected for the pumpkin and acorn cultivars. However, their improvement can be expected to progress at a slower pace than that of the summer squashes for two reasons. First, the summer squashes have greater economic value in the developed countries, where most of the breeding is done. Second, certain cultivars of C. maxima and C. moschata are adapted to areas in which C. pepo is grown and their fruits tend to have superior quality and nutritive value (Culpepper and Moon 1945; Kubicki and Walczak 1976; Whitaker and Davis 1962). Some additional developments in C. pepo will probably result from unpredictable, and seemingly capricious, consumer desires. A case in point is the 'Vegetable Spaghetti' cultivar, a member of the vegetable marrow group. This unusual cultivar has been in commerce in North America since 1936, when it was intro-

440 ECONOMIC BOTANY [VOL. 43 duced by the Burpee Seed Co., and it probably originated in the Far East (T. C. Torrey, pers. comm.). It suddenly gained a surge of popularity in North America around 1980. For 1986, a hybrid cultivar developed in Israel from 'Vegetable Spaghetti' was released (Paris et al. 1985). This new cultivar, named 'Orangetti', has semi-bush habit, intense orange fruit rind and flesh color, and a 15-fold increase in the carotene content of the flesh. Should this cultivar become popular, others like it can be expected to be developed. CONCLUSIONS The edible cultivar groups of C. pepo have existed in their respective recognizable states for a considerable length of time. The pumpkins and scallops were grown in the New World prior to the Discovery in 1492. Pumpkins entered recorded history no later than 1542 and scallops by 1554. Records of pumpkins and scallops have appeared continually since. The acorns were depicted as early as 1591 and again in 1757. The acorns were described in Europe in the 1800s, and a color illustration appeared in France in 1860. No North American record of them was found that predated 1913. No description of the crooknecks has been found in early botanical writings. While the earliest indisputable record is from 1828, with continual recordings thereafter, the group is probably considerably older. The straightnecks are probably the most recently constituted cultivar group, with a history dating back only as far as 1896. The apparent forerunners of the vegetable marrows were illustrated as early as 1591 and again in the 1600s. No additional record concerning this group was found that predated 1800, but after that date the group is mentioned in the literature continually. A forerunner of the cocozelles and zucchinis was described in the early 1600s. The cocozelles have been recorded continually since 1847 and the zucchinis since 1856. The majority of modern cultivars of each of the groups appears to have a common ancestry. The pumpkins, scallops, and crooknecks probably first originated in North America from wild ancestors. The straightnecks, acorns, vegetable marrows, cocozelles, and zucchinis appear to be their derivatives. Of these five derived groups, all but the straightnecks appear to have originated in Europe. Most of the eight groups have been and continue to be of local or regional importance. Only the pumpkins and zucchinis can be considered to be of worldwide distribution and importance. In contrast to the pumpkins, the zucchinis are present mainly in the developed countries and their rise to prominence has been recent. ACKNOWLEDGMENTS I express my sincere appreciation to Ruthie Brumbaugh of the Interiibrary Loan Office, Hume Library, University of Florida, Gainesville; to Dan Nadel of the Hazera' Seed Co., Mivhor, Israel; to the staff of the New York Botanical Garden Library, for obtaining or providing reference material to

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