The Paleobiolinguistics of Domesticated Manioc (Manihot esculenta) 61  Research Communica on  eastern Acre, Brazil, and in immediately adjacent areas in lowland Bolivia (Schaal et al. 2006). Populations of ssp. flabellifolia in northern South America are shown by Léotard et al. (2009) not to be involved in domesti- cation, and other species in Mesoamerica and South America are ruled out by Duputié et al. (2011). Manioc can be lethally toxic if not prepared correctly. The presence of cyanogenic glucosides in the roots varies from less than 10 to more than 500 mg hydrogen cyanide (HCN) per kilo on a fresh weight basis; varieties with less than 100 mg/kg are considered sweet, and those with more are considered bitter (McKey et al. 2010) and require processing for consumption. While sweet and bitter varieties are difficult to distinguish morphologically, farmers typically are able to segregate bitter from sweet in their production systems (McKey and Beckerman 1993). Initial human selection probably favored the development of sweet varieties, which are more widely distributed in South America and through Central America to Mexico than bitter varieties (Arroyo-Kalin 2010). When food production systems started supplying significant proportions of human diets between 4000 and 3000 BP (Piperno and Pearsall 1998), bitter varieties were selected for toxicity to protect against pests and to provide higher yields (Arroyo-Kalin 2010). Bitter varieties are most common in Central and Eastern Amazonia and the Guianas, with sweet varieties being more prevalent in Paleobiolinguistics (PBL) employs the comparative method of historical linguistics to reconstruct the biodiversity known to human groups of the remote, unrecorded past (Brown et al. 2013a; Brown et al. 2013b).1 Comparison of words for biological taxa from languages of the same language family facilitates reconstruction of the biological vocabulary of the family’s ancient proto-language. This study uses PBL to establish where and when domesticated manioc (Manihot esculenta Crantz, Euphorbiaceae) developed significance for different prehistoric Native American groups. This entails mapping in both time and geographic space proto-languages for which words for manioc reconstruct. The approximately 98 species of Manihot are all native to the New World. There are two centers of diversity of Manihot. One is in Mesoamerica, which current evidence indicates may be the place of origin of the genus, with approximately 17 species; and the other is in Brazil, with approximately 80 species (Duputié et al. 2011). The only domesticated taxon is M. esculenta ssp. esculenta, which is derived from ssp. flabellifolia. This subspecies is widely distributed in tropical South America in seasonal open forests both north and south of the Amazon basin, and extending into the Brazilian savannas (Allem 2002). Work during the last decade establishes that ssp. esculenta was brought into domestication somewhere in what are today northwestern Mato Grosso, Rondônia, and The Paleobiolinguis cs of Domes cated Manioc (Manihot esculenta) Cecil H. Brown1*, Charles R. Clement2, Pa ence Epps3, Eike Luedeling4, and Søren Wichmann5  Author address: 1Northern Illinois University, 1700 Scenic Highway, #601, Pensacola, FL, 32503‐6634, USA, 2Ins tuto  Nacional de Pesquisas da Amazônia, Manaus, AM, Brazil, 3 University of Texas at Aus n, Aus n, TX, USA, 4World Agroforest‐ ry Centre (ICRAF), Nairobi, Kenya, 5Max Planck Ins tute for Evolu onary Anthropology, Leipzig, Germany.  *Corresponding author: brown.cecil@yahoo.com   Received: January 21, 2013  Volume: 4:61‐70  Published: June 26, 2013  © 2013 Society of Ethnobiology  Abstract: Paleobiolinguis cs  is  used  to  iden fy  on maps  where  and  when manioc  (Manihot  esculenta)  developed  im‐ portance for different prehistoric groups of Na ve Americans. This  informa on  indicates, among other things, that signifi‐ cant interest in manioc developed at least a millennium before a village‐farming way of life became widespread in the New  World.   Key Words: Archaeobotany, crop origins, historical linguis cs, Na ve American Indians, paleobiolinguis cs, plant domes ca‐ on, plant gene cs    mailto:brown.cecil@yahoo.com� 62  Research Communica on  Years  Before  Present      Proto‐Language  Proto‐Word for  Manioc (NR = Not  Reconstructable)  Homeland Center  Geographic   Coordinates      Family Affilia on    Proto‐Word  Source  6591  Otomanguean  *ya  18, ‐96.92  Otomanguean  1  5498  Popolocan‐ Zapotecan  *ya  17.17, ‐96.17  Otomanguean  Authors  4274  Totozoquean  *pisi:  19.92, ‐97.42  Totozoquean  2  3149  Zapotecan  *ko: yaka  17.17, ‐96.17  Otomanguean  Authors  3036  Popolocan  *ya  18, ‐96.92  Otomanguean  Authors  2445  Chiapanec‐Mangue  *yáɁ  17.07, ‐92.73  Otomanguean  3  2220  Mayan  *tz’ihn  15.42, ‐91.83  Mayan  4  1935  Chinantecan  *ɁmaL  17.92, ‐96.5  Otomanguean  5  1676  Zapotec  ko yaka  17.17, ‐96.17  Otomanguean  Authors  1649  Quichean‐Mamean  *tz’iin  15.42, ‐91.83  Mayan  4  1596  Mixe‐Zoquean  *pisi  17.22, ‐96.03  Totozoquean  6  1520  General Aztec  *kwaw‐kamoʔ  18.35, ‐99.83  Uto‐Aztecan  Authors  1492  Greater Mamean  *tz’iin  15.42, ‐91.83  Mayan  4  1435  Totonacan  *qoqšqḛwḭ  19.92, ‐97.42  Totozoquean  7  1432  Cholan‐Tzeltalan  *tz’ihn  16.83, ‐92.83  Mayan  4  1225  Kanjobalan‐Chujean  *tz’iin  15.83, ‐91.83  Mayan  4  1148  Cholan  *tz’ihn  14.81, ‐89.38  Mayan  4  1058  Chujean  *tz’in  15.92, ‐91.58  Mayan  4  997  Cha no  *ko: yaka  16.25, ‐97.38  Otomanguean  Authors  981  Greater Quichean  *tz’iin  14.78, ‐91.5  Mayan  4  900  Mixe  *kuhy‐piši  17.02, ‐96.07  Totozoquean  6  802  Kanjobalan  *tz’iin  15.83, ‐91.83  Mayan  4  790  Yucatecan  *tz’iin  20, ‐89  Mayan  4  787  Zoque  *pisi  16.9, ‐94.68  Totozoquean  6  511  Tzeltalan  *tz’in  16.83, ‐92.83  Mayan  4  Table 1. Manioc‐term reconstruc on for proto‐languages of Mesoamerica (Southern Mexico and Northern Central  America).  Proto‐Word Source:  1. Kaufman 1990   2. Brown et al. 2011  3. Rensch 1976  4. Brown and Wichmann 2004  5. Rensch 1989  6. Wichmann 1995    7. David Beck, pers. com.  the basin’s headwaters (McKey and Beckerman 1993). Bitter varieties were also once common along coastal Brazil, but now are rare. Isendahl (2011) reviews the archaeobotanical evidence for cultivated manioc, with oldest dates of approximately 8000 BP for remains from the Pacific coast of Peru and low-elevations of the Colombian Andes, 5000 BP for lowland Amazonian Colombia, 7000 BP for Panama, and 6500 BP for Mexico. Arroyo-Kalin (2010) cautions that early dates for manioc may not always be M. esculenta, since the genus is widely distributed and other species may have been brought into domestication and later abandoned with the arrival of modern manioc. However, given that other Manihot species are absent from coastal Peru and the Colombian Andes (Duputié et al. 2011), the 63  Research Communica on  earliest dates in these regions are likely to represent the introduction of M. esculenta from its center of domestication in southwestern Amazonia. Manioc-term reconstructions are presented for proto-languages of two major regions of the New World: (1) Southern Mexico and Northern Central America (henceforth Mesoamerica) (Table 1); (2) Southern Central America and South America (Table 2). North America is not included in this analysis because manioc was a historical introduction there. The tables list major proto-languages of the Americas widely regarded by historical linguists as demonstrated. Some major proto-languages are not included because lexical information from daughter languages is not sufficiently available for drawing either positive or negative conclusions about manioc- term reconstruction. Unfortunately, words for sweet and bitter varieties are insufficiently recorded in sources to permit a reconstructed distinction. In addition to identifying proto-languages with manioc terms and the terms themselves, the tables report proto-languages for which these terms are “not reconstructable” (NR). NR is a designation used when terms for manioc are present in all or most languages of a family, but, nonetheless, are not cognate and, hence, do not attest to a manioc term in their shared ancestral language. NR, then, never indicates that a term does not reconstruct because of missing data. Dates for proto-languages presented in the tables are intended to be the latest dates at which these languages were spoken just before breaking up into daughter languages. These are calculated through use of Automated Similarity Judgment Program (ASJP) chronology, a computational dating approach based on the lexical similarity of languages (Holman et al. 2011).2 Possible geographic coordinates for proto- language homeland centers given in the tables are produced through automation using an algorithm for identifying the maximum lexical diversity within a language family (Wichmann et al. 2010). The geo- graphic center of lexical diversity of a family is assumed to correlate with where the family’s proto- language was spoken. Tables also give a linguistic Figure 1. Manioc‐term reconstruc on informa on from Table 1 plo ed on map of Mesoamerica.   64  Research Communica on  Table 2. Manioc‐term reconstruc on for proto‐languages of Southern Central America and South America.  Years  Before  Present      Proto‐Language  Proto‐Word for  Manioc (NR = Not  Reconstructable)  Homeland Center  Geographic       Coordinates      Family Affilia on      Proto‐Word Source  7266  Macro‐Ge  NR  ‐11.3, ‐53  Macro‐Ge     4701  Mataco‐Guaykuru  NR  ‐22.5, ‐62.58  Mataco‐ Guaykuru     4461  Southern Arawakan  *kaniri  ‐10.33, ‐74.33  Arawakan  Authors  4400  Chibchan  *Ɂik, *ike  9.75, ‐83.42  Chibchan  1, 2  4134  Arawakan  *kani[tʰɨ]  1, ‐69.17  Arawakan  3  4085  N Arawakan  *kani[ ]  1, ‐69.17  Arawakan  Authors  3943  Panoan‐Tacanan  NR  ‐7.5, ‐75  Panoan‐Tacanan     3585  Tupi  *mani  ‐8, ‐62  Tupi  4  3518  Caribbean N Arawakan  *kani  12, ‐72  Arawakan  Authors  3310  Salivan  NR  5, ‐67  Salivan     3241  Barbacoan  *ku‐  0.67, ‐79  Barbacoan  Authors  3178  Zaparoan  *muɾɨha  ‐3.25, ‐74  Zaparoan  5  3124  Nadahup  NR  0, ‐69  Nadahup     3023  Ge  *kwyr  ‐15, ‐52.5  Macro‐Ge  6  2927  Witoto‐Ocaina  *hõɁ   ‐2.75, ‐71.75  Witoto‐Ocaina‐ Nonuya  7  2909  Guaykuruan  NR  ‐26.5, ‐59  Mataco‐ Guaykuru    2857  Witoto‐Ocaina‐Nonuya  *hoɁ t  ‐1.25, ‐72.5  Witoto‐Ocaina‐ Nonuya  8  2807  Nambiquaran  *(wv3)lin’3  ‐12.75, ‐59.17  Nambiquaran  9  2774  Misumalpan  NR  13, ‐84.5  Misumalpan     2731  Talamancan  *ik  9.75, ‐83.42  Chibchan  Authors  2699  Tucanoan  *kɨi  0.33, ‐70.25  Tucanoan  10  2593  Inland N Arawakan  *kainhi, *kap(w)a‐ li, *mulhui  1, ‐69.17  Arawakan  11  2503  Venezuelan Cariban  *kɨčere, *amaka  6.5, ‐66  Cariban  Authors  2433  Southern Guaykuruan  *piyok  ‐26.5, ‐59  Mataco‐ Guaykuru  Authors  2414  North Barbacoan  *ku‐  1.5, ‐78.25  Barbacoan  Authors  2412  Cariban  *kï(č/t)ere, *wïj(u)  10.17, ‐72.75  Cariban   12  2404  Matacoan  NR  ‐22.5, ‐62.58  Mataco‐ Guaykuru     2271  Boran  *pɨɨkkaá,  *paikoómɨ  ‐2.17, ‐72.33  Boran  7,  Authors  2219  Purus  *kanшrш  ‐12.5, ‐69.33  Arawakan  Authors  2156  Western Tucanoan  *kɨi  ‐2.83, ‐72.5  Tucanoan  Authors  1931  Chapacuran  *Ɂakop  ‐13.43, ‐63.17  Chapacuran  13  1853  Panoan  *Ɂatsa  ‐7.5, ‐75  Panoan‐Tacanan  Authors  1853  Pekodian  *u  ‐14, ‐55  Cariban  Authors  1850  Tupari  *mani  ‐12.5, ‐62.5  Tupi  Authors  (con nued on next page)  65  Research Communica on  1821  Southern Ge  *kɯbɛ{d}, *kagre  ‐26, ‐52  Macro‐Ge  14  1780  Mascoian  *seppo  ‐23.2, ‐58  Mascoian  Authors  1764  Arauan  *po'a, *ximeka,  *kojo  ‐6, ‐70.5  Arauan  15  1717  Quechuan  NR  0.33, ‐78  Quechuan  16  1712  Monde  NR  ‐10.97, ‐61.13  Tupi     1634  Mainline Panoan  *Ɂatsa  ‐7.5, ‐75  Panoan‐Tacanan  17  1607  Jabu   *murε  ‐12.25, ‐62.25  Macro‐Ge  18, 19  1590  Tacanan  *kwawe  ‐13.33, ‐66.5  Panoan‐Tacanan  20  1569  Harakmbet  *táare  ‐12.5, ‐70.5  Harakmbet  21  1550  Tupi‐Guarani  *mani  ‐8, ‐62  Tupi  5  1519  Kampan  *kaniri  ‐10.33, ‐74.33  Arawakan  Authors  1480  Munduruku  *masɨk  ‐7, ‐58  Tupi  Authors  1418  Cayapa‐Colorado  *kuhcu  0.67, ‐79  Barbacoan  22  1402  Guianan Cariban  *kičere, *wɨi  3.25, ‐55.75  Cariban  Authors  1395  Cabecar‐Bribri  *ali  9.42, ‐83  Chibchan  Authors  1335  Kakua‐Nukak  *tũj  0.88, ‐69.56  Kakua‐Nukak  Authors  1319  Yanomam  *naši  3.5, ‐62.83  Yanomam  Authors  1291  Guahiban  *newa, *bawá  6.5, ‐71.33  Guahiban  23,  Authors  1262  Timbira  *kwɨr  ‐5.25, ‐46  Macro‐Ge  18  1241  Eastern Tucanoan  *kɨi  0.33, ‐70.25  Tucanoan  Authors  1185  Kawapanan  *kiɁ  ‐5.5, ‐77  Kawapanan  24  1169  Pemongan  *kɨsera  4, ‐60  Cariban  Authors  992  Taranoan  *wɨi  1, ‐73  Cariban  25  974  Quechua II  *rumu  0.33, ‐78  Quechuan  16  678  Jivaroan  *máma  ‐2.5, ‐78  Jivaroan  26  644  Guaymi  *ho, *hi  8.67, ‐82  Chibchan  Authors  419  Coconucan  *lo  2.5, ‐76.5  Barbacoan  Authors  414  Witoto Proper  *hu , *maika(hi)  ‐1, ‐73.5  Witoto‐Ocaina‐ Nonuya  Authors  389  Mayoruna Panoan  *Ɂatsa  ‐4.42, ‐70.25  Panoan‐Tacanan  Authors  Years  Before  Present      Proto‐Language  Proto‐Word for  Manioc (NR = Not  Reconstructable) Homeland Center  Geographic       Coordinates      Family Affilia on      Proto‐Word Source  Proto‐Word Source:  1. Constenla 1981   2. Constenla 1990   3. Payne 1991  4. Rodrigues 2010   5. Lev Michael, pers. com.  6. Davis 1966   7. Aschmann 1993  8. Echeverri and Seifart 2011  9. Price 1978  10. Chacon n.d.  11. Ramirez 2001  12. Sergio Meira, pers. com.  13. Angenot‐de Lima 1997   14. Jolkesky 2010   15. Dixon 2004  16. Willem Adelaar, pers. com.  17. Shell 2008  (con nued from previous page)  18. Eduardo Ribeiro, pers. com  19. van der Voort 2007  20. Girard 1971  21. Ma eson 1972   22. Moore 1962  23. Chris an and Ma eson 1972   24. Pilar Valenzuela, pers. com.  25. Meira 2000  26. Payne 1981   66  Research Communica on  family affiliation for each proto-language. The information reported in Tables 1 and 2 is plotted on maps of Figures 1, 2 and 3 to give a visual perspective on both the chronological and geographic distribu- tions of reconstructed manioc terms. Figure 3 is an enlargement of a highly congested area of the map of Figure 2. As noted above, manioc was originally domesti- cated somewhere in southwestern Amazonia (Schaal et al. 2006). This area includes the presumed Tupi homeland (Noelli 1998; Rodrigues 1964; Walker et al. 2012; and see Figure 2). Nevertheless, the ASJP date for Proto-Tupi is somewhat more recent (3585 BP, Table 2) than dates for other South American groups, especially Arawakan (4461-4085 BP, Table 2). The Arawakan homeland is near the 5000 BP-manioc remains reported from lowland Colombian Amazonia (Isendahl 2011) and, thus, appears to reflect the dispersal of the crop from its center of domestication. From PBL analysis, there is no direct indication that speakers of any particular South American proto- language were responsible for manioc domestication. While archaeological evidence attests to the presence of manioc in Central America as early as 7000 BP (Isendahl 2011), the earliest date for a proto- language in the area having a manioc term, i.e., Chibchan, is 4400 BP (Table 2). This 2600-year discrepancy may be more apparent than real consider- ing, for example, that ASJP dates are minimum dates for proto-languages, i.e., latest dates at which ancestral languages were still spoken. The date for Oto- manguean, 6591 BP (Table 1), of northern Mesoamer- ica, corresponds well with the archaeological date of 6500 BP for the earliest remains of manioc in the area Figure 2. Manioc‐term  reconstruc on  informa on  from Table 2 plo ed on map of Southern Central America and South  America.   67  Research Communica on  (Isendahl 2011). Notably, manioc reconstructs for all proto-languages of Mesoamerica (Figure 1), while this is not so for those of southern Central America and South America (Figure 2). In general, proto-languages for which manioc terms reconstruct are broadly distributed through lowland South America and Mesoamerica, reflecting the crop’s dispersal throughout the Neotropics from its origin in southwestern Amazonia. ASJP dates for these proto-languages, like archaeobotanical ones, indicate that domestication and dispersal of the crop occurred before the general development of a village- farming way of life in the New World from 4000 to 3000 BP (Piperno and Pearsall 1998; Smith 1992). Acknowledgements We thank Jan Salick, Doyle McKey, Manuel Arroyo- Kalin, Christian Isendahl, two anonymous reviewers, and the editors for useful suggestions for improving this study. We are also grateful to Willem Adelaar, David Beck, Sergio Meira, Lev Michael, Eduardo Ribeiro, and Pilar Valenzuela for supplying linguistic data. Declarations Permissions: Not applicable. Sources of funding: Epps’ work on this project was supported by the National Science Foundation (HSD0902114). Conflicts of interest: None declared. Figure 3. Enlargement of congested region (northwestern South America and adjacent areas) of map of Figure 2.   68  Research Communica on  References Cited Allem, A. C. 2002. The Origins and Taxonomy of Cassava. In Cassava, Biology, Production and Utilization, edited by R. J. Hillocks, J. M. Thresh, and A. C. Belloti, pp. 1-16. CAB International, Wallingford, UK. Angenot de Lima, M. G. 1997. Fonotática e Fonologi do Lexema Protochapakura. Unpublished Master’s Thesis, Universidade Federal de Rondônia. Arroyo-Kalin, M. 2010. The Amazonian Formative: Crop Domestication and Anthropogenic Soils. Diversity 2:473-504. Aschmann, R. P. 1993. Proto Witotoan. 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PLoS ONE 7(4): e35025. doi:10.1371/ journal.pone.0035025 Wichmann, S. 1995. The Relationship among the Mixe- Zoquean Languages of Mexico. University of Utah Press, Salt Lake City. Wichmann, S., A. Müller, and V. Velupillai. 2010. Homelands of the World’s Language Families: A Quantitative Approach. Diachronica 27:247–276. Biosketches Cecil H. Brown is a linguis c anthropologist with  interests in ethnobiology, historical linguis cs, and  Na ve American languages.  Charles R. Clement is a gene cist studying the origin and  domes ca on of na ve Amazonian crops, and the  ethnobotany associated with anthropogenic soils and  70  Research Communica on  other domes cated landscapes.  Pa ence Epps is a linguist whose work inves gates  lowland South American languages from historical,  typological, and descrip ve perspec ves.  Eike Luedeling is an agricultural scien st mainly  concerned with projec on of climate change impacts on  agricultural and natural ecosystems and with the  development of appropriate adapta on strategies.  Søren Wichmann specializes in quan ta ve methods in  historical linguis cs and Mesoamerican languages. He is  General Editor of the journal Language Dynamics and  Change.  Notes 1This is the second PBL study published in Ethnobiolo- gy Letters, the first treating chili pepper (Brown et al. 2013a). Several others are projected. The method and theory of PBL is summarized in Brown et al. (2013a) and discussed in substantial detail in several preceding papers (Brown 2006a,b; Brown 2010; Brown et al. 2013b). The current paper and others that follow in Ethnobiology Letters will not repeat these discussions. 2Occasionally, an ASJP date for a proto-language may be older than a date for its own parent language. For example, Proto-Southern Arawakan (4461 BP) has an ASJP date older than that for Proto-Arawakan (4134 BP). This sometimes occurs in ASJP chronology when a language group’s breakup is closely followed in time by the breakup of its immediate subgroup. 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