Caldasia 23(1): 43-76 TOW ARD A CLADISTIC MODEL FOR THE CARIBBEAN SUBREGION: DELIMITATION OF AREAS OF ENDEMISM

JUAN J. MORRONE Museo de Zoología, Facultad de Ciencias, UNAM, Apartado 70-399, 04510 México D.F., México.

RESUMEN Se propone un nuevo esquema biogeográfico para la subregión Caribeña, de acuer- do con el cual se reconocen las siguientes 29 provincias: Sierra Madre Occidental (oeste de México: estados de Chihuahua, Durango, Zacatecas, Sonora, Sinaloa, Nayarit y Jalisco, por encima de los 1000 m), Sierra Madre Oriental (este de México: estados de San Luis Potosí, Coahuila, Hidalgo, Nuevo León y Querétaro, por encima de los 1500 m), Eje Volcánico Transmexicano (centro de México: estados de Guanajuato, México, Distrito Federal, Jalisco, Michoacán, Puebla, Oaxaca, Tlaxcala y Veracruz), Depresión del Balsas (centro de México: estados de Guerrero, México, Jalisco, Michoacán, Morelos, Oaxaca y Puebla, por debajo de los 2000 m), Sierra Madre del Sur (centro sur de México, desde el sur de Michoacán hasta Guerrero y Oaxaca, y parte de Puebla, por encima de los 1000 m), Golfo de México (costa del Golfo de México, en el este de México, Belice y norte de Guatemala), Península de Yucatán (Península de Yucatán, en los estados mexicanos de Campeche, Yucatán y Quintana Roo, por debajo de los 200 m), Costa Pacífica Mexicana (oeste de México, en la costa pacífica de los estados de Sinaloa, Nayarit, Colima, Jalisco, Michoacán, Guerrero, Oaxaca y Chiapas), Chiapas (México, Guatemala y Nicaragua, básicamen- te corresponde a la Sierra Madre de Chiapas, desde 500 a 2000 m), Oriente de Amé- rica Central (este de América Central, desde Guatemala hasta Panamá), Oeste del Istmo de Panamá (oeste de América Central, desde Costa Rica hasta el oeste de Panamá), Bahamas (archipiélago de las Bahamas, comprende las islas de Abaco- Grand Bahama, Andros-Bimini, Cat, Crooked-Mayaguana, Exumas, Inaguas, Long- Ragged Island Range, Mona, New Providence-Eleutheras, San Salvador-Rum Cay, St. Eustatius, St. Kitts, St. Lucia, St. Martin, St. Vincent, y Turks y Caicos), Cuba (isla de Cuba), Islas Caimán (archipiélago que incluye las islas Grand Cayman, Little Cayman y Cayman Brac), Jamaica (isla de Jamaica), Española (República Dominica- na y Haití), Puerto Rico (isla de Puerto Rico), Antillas Menores (archipiélagos de las Antillas Menores y las Islas Vírgenes), Maracaibo (norte de Colombia y noroeste de ), Costa Venezolana (norte de Venezuela y Colombia; también incluye las islas de Aruba, Curacao y Bonaire), Trinidad y Tobago (islas de Trinidad y Tobago), Chocó (costa pacífica del norte de Ecuador, Colombia y Panamá), Cauca (Colombia y Ecuador), Magdalena (oeste de Venezuela y noreste de Colombia), Llanos Venezo- lanos (gran parte de Venezuela y noreste de Colombia), Ecuador Occidental (oeste de Ecuador y sudoeste de Colombia), Ecuador Seco (oeste de Ecuador), Islas Galápagos (archipiélago de Colón, en el océano Pacífico, 950 km al oeste de la costa ecuatoriana) y Tumbes-Piura (sur de Ecuador y norte de Perú). Cada una de las Toward a cladistic model Ior thc Caribbcan subrcgion

provincias representa una unidad biogeográfica natural, definida por la coinciden- cia de las distribuciones de varios taxones animales y vegetales.

Palabras clave. Caribe, Neotrópicos, biogeografía, cndemisrno.

A BSTRACT A new biogeographic scherne for the Caribbean subregion is proposed herein, where the following 29 provinces are recognized: Sierra Madre Occidental (western Mexico: states of Chihuahua, Durango, Zacatecas, Sonora, Sinaloa, Nayarit, and Jalisco, above 1,000 m altitude), Sierra Madre Oriental (eastern Mexico: states ofSan Luis Potosí, Coahuila, Hidalgo, Nuevo León, and Querétaro, above 1,500 m altitud e), Transmexican Volcanic Belt (central Mexico: states ofGuanajuato, Mexico, Distrito Federal, Jalisco, Michoacán, Puebla, Oaxaca, Tlaxcala, and Veracruz), Balsas Basin (central Mexico: states ofGuerrero, Mexico, Jalisco, Michoacán, Morelos, Oaxaca, and Puebla, below 2,000 m altitude), Sierra Madre del Sur (south central Mexico, from southern Michoacán to Guerrero and Oaxaca, and part of Pucbla, abo ve 1,000 m altitude), Mexican Gulf (coast ofthe Mexican Gulf, in eastern Mexico, Belize, and northern Guatemala), Yucatán peninsula (Yucatán península, in the Mexican states of Carnpeche, Yucatán, and Quintana Roo, below 200 m altitude), Mexican Pacific Coast (western Mexico, in the Pacific coast ofthe states ofSinaloa, Nayarit, Colima, Jalisco, Michoacán, Guerrero, Oaxaca, and Chiapas), Chiapas (Mexico, Guatemala, and Nicaragua, basically corresponding to the Sierra Madre de Chiapas, from 500 to 2,000 m altitude), Eastern Central Arnerica (eastern Central America, frorn Guatemala to Panarna), Western Panamanian Isthrnus (western Central America, from Costa Rica to western Panama), Bahama (Bahama archipelago, comprising the islands of Abaco-Grand Bahama, Andros-Birnini, Cat, Crooked-Mayaguana, Exurnas, Inaguas, Long-Ragged Island Range, Mona, New Providence-Eleutheras, San Salvador-Rum Cay, St. Eustatius, St. Kitts, St. Lucia, St. Martin, St. Vincent, and Turks and Caicos), Cuba (island ofCuba), Cayman islands (archipelago including Grand Cayman, Little Cayrnan, and Cayrnan Brac), Jamaica (island ofJamaica), Hispaniola (Dominican Republic and Haiti), Puerto Rico (island ofPuerto Rico), Lesser Antilles (archipelagos ofthe Lesser Antilles and the Virgin islands), Maracaibo (northern Colombia and northwestern Venezuela), Coastal Venezuela (northern Venezuela and Colombia; also includes the islands of Aruba, Curacao, and Bonaire), Trinidad and Tobago (islands ofTrinidad and Tobago), Chocó (Pacific coast ofnorthern Ecuador, Colom- bia, and Panarna), Cauca (Colombia and Ecuador), Magdalena (western Venezuela and northeastern Colombia), Venezuelan Llanos (great part of Venezuela and northeastern Colombia), Western Ecuador (western Ecuador and southwestern Colombia), Dry Ecuador (western Ecuador), Galapagos islands (Colón archipelago, in the Pacific Ocean, 950 km west frorn the coast of Ecuador), and Tumbes-Piura (southern Ecuador and northern Peru). Each ofthe provinces represents a natural biogeographic unit, herein defined by the coincidence ofthe distributional ranges of several and taxa.

Key Words. Caribbean, Neotropics, biogeography, endemism.

44 Juan J. Morronc INTRODUCTION restricted to portions of the subregion (Ryan, 1963; Rauchenberger, 1988; Aguilar el al, Disjunctions in the distributional ranges of 1997; Morrone, 1999; Morrone el al., 1999). several plant and animal taxa in continental My objective herein is to integrate these and insular areas bordering the Caribbean sea previous studies, in order to provide a com- are among the most familiar patterns known plete regionalization of the Caribbean to biogeographers (Croizat, 1976; Rosen, 1976, subregion. The provinces recognized in this 1985; Halff1:er, 1978, 1987; MacFadden, 1980; scheme are intended to constitute areas of Pregill, 1981; Savage, 1982; Liehbherr, 1988a; endernism, which could be considered as units Rauchcnberger, 1988; Williams, 1989; Zunino, in a future cladistic biogeographic analysis of 1990; Hedges el at., 1992; Page & Lydeard, the subregion. 1994; Lydeard el al, 1995; Crother & Guyer, 1996; Hedges, 1996a, b; Llorentc, 1996; MATERIAL AND METHODS Escalante el al., 1998). Sorne of the published stu d ies ha ve foc u sed on postu lati ng Dislributional data for this study were hypotheses to explain the origin 01' disjunct obtained frorn the literature. In addition, patterns, either frorn the dispersalist or biogeographic schernes proposed by previous vicariancc viewpoints. The formal recognition authors were compiled and the units of t h e wh o lc area from a descriptive recognized by thern were cornpared under a perspective and of the arcas of endern ism track approach, in order to test whether they within it, however, has been usually neglected. represented natural biogeographie units. The It is increasingly recogn ized that the dispersal/ track approach basically eonsists in detecting vicariance opposition -recently exposed in generalized tracks, which indicate the relation to the Caribbean by the interchange preexistence ofancestral biotas. A generalized between Hcdges el al (1992), Page & Lydeard traek is equated herein to a natural (1994), Crother & Guyer (1996), and Hcdges biogeographic unit or an area of endernism. (1996a)- should be abandoned. As clcarly When different generalised tracks intersect stated by Page & Lydeard (1994), a more fun- in a given area, they determine a node, which damental biogeographic question is to search indicates that different ancestral biotic or for congruent patterns among area geological fragments interrelate, as a cladograms, which would suggest a common consequence ofterrain collision, docking or cause, be it vicariance, dispcrsal or (rnost suturing, thus constituting cornposite biotas probably) a cornbination of both processes. or areas. A node is considered herein to be A basic premiso ofa cladistic biogeographic located in the boundaries among different na- analysis is the recognition of both the area tural areas. (For details of the track under study and the units within it (Morrone, rncthodology see Morrone & Crisci, 1995 and 1994). Lumping of largo arcas into poorly Craw el al, 1999.) dcfined arca, e.g., "nuclear Central América" For the subregion and each province, a brief (Savage, 1982; Rauchenberger, 1988), "lower characterization is provided, sorne of the Central América" (Savage, 1982), "rnainland" different names received by the areas are (Hedges el al., 1992), "West ludies" (Hedges Iistcd, and the taxa considered to support thern el al, 1992), "Central America" (Page & are detailed. Synonymies should be Lydeard, 1994), will hardly reflect natural arcas. considered as approximate, as a complete Although sorne studics have defined areas of coincidence between the different names is cndernism in the Caribbean, they have been almost never achieved. In the taxonornic lists,

45 Toward a c1adistic modc\ for the Caribbean subregion acronyms of the authors are detailed in 71), Central American province (Fittkau, 1969: appendix 1. 642), 1ncasic province (Fittkau, 1969: 642), Caribbean subregion Mountain Mesoamerican province (Cabrera The Caribbean subregion is the most septen- & Willink, 1973: 32), Caribbean domain (Ca- trional ofthe Neotropical region, comprising brera & Willínk, 1973: 32), Guajira province most of Mexico, Central America, the West (Cabrera & Willink, 1973: 46), Pacific province lndies, and northwestern South America (Cabrera & Willink, 1973: 52), Venezuelan (Morrone, 1999). According to the new province (Cabrera & Willink, 1973: 56), biogeographic classification presented herein, Orinoco- Venezuelan domain (Ringuelet, 1975: the Caribbean subregion comprises most1y 107), Pacific domain (Ringuelet, 1975: 107), areas ringing the Caribbean sea: part of Mountain Mesoamerican region (Rzedowski, Mesoamerica to the west, southern South 1978: 101), Caribbean region (Rzedowski, 1978: America to the south, the Lesser Antilles to 107; Takhtajan, 1986: 251; Rangel et al., 1995c: the east, and the Greater Antilles (Cuba, Ja- 21; Liebherr, 1997: 89), West Indian region maica, Hispaniola, and Puerto Rico) to the (Nichols, 1988: 71), Orinoco endemic region north. Other provinces are placed in areas (Vari, 1992a: 13), Westem endemicregion (Vari, ringing the central Pacific ocean, from Mexico 1992a: 14), Caribbeo-Mexican region (Rivas- to Ecuador, as well as the Galapagos Islands. Martínez & Navarro, 1994: map), Colombian- This proposal assigns the Caribbean sea and Mesoamerican region (Rivas-Martínez & Na- the central Pacific ocean as the baselines of varro, 1994: rnap), Venezuelan region (Rivas- the Caribbean subregion. Baselines (Croizat, Martínez & Navarro, 1994: map), Caribbean 1952) are working hypotheses by which sets area (Coscarón & Coscarón-Arias, 1995: 726), of disjunctions can be historically linked, Pacific area (Coscarón & Coscaron-Arias, providing a locus around which to rationalize 1995: 726; Morrone & Coscarón, 1996: 2), explanations of geographic distribution (Craw Northern Andes bioregion (Dinerstein et al., et al., 1999). As stated by Craw & Page (J 988) 1995: map), Middle America (Campbell, 1999: and Parenti (1991), the major patterns of 112), and Caribbean subregion (Morrone, geographic distribution cross ocean basins, 1999: 2; Morroneetal., 1999: 511). thus showing that the c\assical realms do not Taxa. FILICOPHYT A. Filicopsida. Filicales. represent natural areas. In this respect, the Pteridaceae: Pellaea ovata (TR). CONI- Caribbean subregion as herein redefined, FEROPHYT A. Coniferopsidae. Coniferales. seems to be a complex area, with biotic Podocarpaceae: Podocarpus guatemalensis affinities across both the Pacific and (TO). MAGNOLlOPHYT A. Liliopsida. Caribbean. The available c\adistic biogeo- Orchidales. Burmanniaceae: Burmanniaflava, graphic analyses (Rosen, 1976; MacFadden, B. kalbreyeri, Gymnosiphon panamensis, 1980; Savage, 1982; Zunino, 1990; Page & Thismia panamensis (MA). Poales. Poaceae: Lydeard, 1994; Crother & Guyer, 1996) suffer Gouninia virgata (OR). Magnoliopsida. from the problem that they only dealt with a Aristo loch iales. A risto loch iaceae: portion of the Caribbean subregion. 1 hope Aristolochia anguicida, A. grandiflora, A. the present scheme will encourage others to inflata, A. odoratissim a, A. ringens, A. undertake a c\adistic biogeographic ana1ysis tonduzii (GO). Asterales. Asteraceae: ofthis subregion as a whole. Montanoa (FU). Fabales. Fabaceae: Synonyrns. North Andean zone (Shannon, Crotalaria maypurensis (BE). Rosales. 1927: 3), Incasic district(Cabrera& Yepes, 1940: Connaraceae: Cnestidium, Con narus 16), Antillean subregion (Rapoport, 1968: 71), panamensis (FO). Urticales. Cecropiaceae: Central American subregion (Rapoport, 1968: Cecropia angustifol ia, C. insignis, C.

46 Juan J. Morrone obtusifolia, C. peltata, C. polyphlebia, C. crocinopterus (MO). Neuroptera. Hemero- telenitida (FR). Violales. Passifloraceae: biidae: Hemer ob iell a, Megalomus Passiflora vitifolia (ES). ANNELIDA. marginatus, M. minor (MN). Orthoptera. Oligochaeta. Megascolecidae: Trigaster(RF). Pyrgomorphidae: Minorissa pustulata (KE). ARTHROPODA. Chilopoda. Geophilomor- Trichoptera. Philopotamidae: Chimarraflinti pha. Ballophilidae: Caritoha-llex, Clavo- (FL); Polycentropidae: Polycentropus philus, Leuconinum (PE); Geophilidae: altmanni, P. insularis species group (FL, HL). Piestophilus, Portoricona, Telocricus (PE); VERTEBRAT A. Actinopterygii. Cyprino- Schendylidae: Algunguis, Bimindyl a, dontiformes. Anablepidae: Anableps (PA); Portoricellus, Schendylops varipictus, S. Cyprinodontidae: Cyprinodon, Girardiini (PA, virgingordae (PE). Crustacea. Decapoda. RA); Poecilidae: Gambusia, Poecilia (RA). Trichodactylidae: Sylviocarcinus piriformis Amphibia. Anura. Bufonidae: Bufo peltoce- (MG, ML, RO). Hexapoda. Blattaria. Blatellidae: phalus species group (HB); Leptodactyl idae: A naplecta mexicana (G U). Coleoptera. Ceratophrys calcarata (L Y). Squamata. Alleculidae: Lobopoda brunneipennis (PK); Gekkonidae: Aristelliger (HB); Iguanidae: Anthribidae: Phaeno-theriopsis (VA); Anolis, Cyclura, Norops (GY, HB); Scindidae: Carabidae: Clivina fasciata, Clivina Mabuya b istriata (HB); Xantusiidae: (Paraclivina), Euphorticus p ubescens ; Lepidophyma (CT). Crocodylia. Alliga- Panagaeus panamensis, P. quadrlsignatus, toridae: Caiman crocodilus fuscus, Platynus spp., Scarites marinus (LE, L1, NI); Crocodylus intermedius (DN, HB). Aves. Cerambycidae: Lagocheirus (TL); Coccine- Coraciiformes. Momotidae: Hylomanes lidae: Sticholodini (GD); Curculionidae: Achia momotula (SI). Craciformes. Cracidae: Ortalis strangulata, Anthonomus marmoratus, A. (SI). Gruiformes. Rallidae: Fulica caribaea subp aralellus, A. unipunctulatus species (MU). Piciformes. Ramphastidae: Selenidera group, Entimus arrogans, Erodiscus spectabilis (CP). Psittaciformes. Psittacidae: attenuatus ; Loncophorus fusiformis, Aratinga nana (SI). Mammalia. Carnivora. Micromyrmex, Neomastix, Prosicoderus Procyonidae: Nasua narica (EM). Chiroptera. scolapax, Pseudoalaocybites, Sicoderus Mormopidae: Mormoops megalophylla (EM); laevigatus (BK, CA, CK, CL, CR, HO, IS, OS, Phyllostomidae: Sturnira ludovici (PC). VN); Erirhinidae: Lissorhoptrus persimilis, Didelphimorphia. Didelphidae: Marmosa Neohydronomus e!egans (OB, OW); Passa- rob insoni, Marmosops fusca tus (EM). lidae: Petrejoides (CE); Sphindidae: Cari- Perissodactyla. Tapiridae: Tapirus bairdii nisphindus (MC); Staphylinidae: Bledius (EM). Primates. Cebidae: Cebus capucinus aequatorialis specres group, B. (EM). Rodentia. Dasyproctidae: Dasyprocta micr oceph alus, B. pu nctat is s im us , punctata (EM); Echimyidae: Diplomys Neohisnius armuellensis (FA, HA, HI). labialis, Hoplomys gymnurus, Proechimys Diptera. Dolichopodidae: Asyn-detus trinitatus species group (PT); Heteromyidae: interruptus, Paraclius desenderi (BS); Heteromys spp. (EM); Muridae: Neusticomys Drosophilidae: Drosophila willistoni and spp., Tylomisspp. (EM); Sciuridae: Microsciurus Zygothrica poeyi species groups (G 1); mimulus, Sciurus granatensis (EM). Xenarthra. Ephydridae: Limnellia huachuca, and Bradypodidae: Cabassouscentralis;C unicinctus, Mimapsilopa cressoni (LZ, MS); Simuliidae: Choloepus hoffmanni (EM); Myrmecophagidae: Simulium metallicum, S. mexicanum, S. Tamandua mexicana (EM). (Psilopelmia) (CB, CC, CO). Hemiptera. Lygaeidae: Ozophora quinquemaculata Sierra Madre Occidental province (Fig. 1.1) complex (SL); Reduviidae: Thymbreus Western Mexico, in the states of Chihuahua,

47 Toward a cladistic modcl for thc Caribbcan subrcgion

Durango, Zacatecas, Sonora, Sinaloa, Nayarit, Magnoliopsida. Ebenales. Styracaceae: Styrax and Jalisco, aboye 1,000 m altitude. argenteus (CG). Juglandales. Juglandaceae: Synonyms. Sierra Madre Occidental province Juglans mollis (AR, RZ). ARTHROPODA. (Rzedowski, 1978: 102; Álvarez el al., 1995: Hexapoda. Coleoptera: Passalidae: 80; Arriaga el al., 1997: 64; Escalante el al., Odoniotaenius zo diacu s, Petrejo ides 1998: 285; Carnpbell, 1999: 1 14; Morrone el laticornis, 1'. nebulosus, P. orizabae, and P. al., 1999: 511) and Sierra Madre Occidental silvaticus (CE, CS); Scarabaeidae: Anopsiostes pine and oak forests ecoregion (Dinerstein el pauliani, Cotinis orientalis, Homoiosternus al., 1995: 102). setosus (DI3, DE, DH); Staphylinidae: Taxa. CONIFEROPHYTA. Coniferopsida. Styngetus deyrollei (NH). . Coniferales. Pinaceae: Pinus engelmanii (AR, Papilionidae: Pterourus palamedes leontis RZ). MAGNOLIOPHYT A. Magnoliopsida. (LL). VERTEBRATA. Squamata. Colubridae: , : madrensis (GE). Rhadinca gaigae (AR). Aves. Galliformes. ARTHROPODA. Hexapoda. Coleoptera: Phasianidae: Dendrortix barbatus (AR). Cleridae: En oclerus m adrensis (RI); Psittaciformes. Psittacidae: Rhynchopsitta Scarabaeidae: Coscinocephalus cribrifrons, t errisi (AR). Strigiformes. Strigidae: C. tepehuanus, Hologymnetis argenteola, Glaucidium sanchezi (AR). Mammalia.

Hom oi osternus beckeri, Onthop h.agus Rodentia. Muridae: Peromyscus (l. aztccus c op or o ides (D8, LH, MF, MR, RD). (SU). VERTEBRAT A. Amphibia. Urodela. Ambystomatidae: Ambystoma rosaceum Transmexican Volcanic Belt province (Fig. (AR). Squamata. Colubridae: Thamnophis 1.3) nigronuchalis (AR); Viperidae: Crotalus Central México, in the states ofGuanajuato, pricei, C. willardi (CV). Aves. Passeriformes. Mexico, Distrito Federal, Jalisco, Michoacán, Corvidae: Cy anoc or ax dickeyi (AR). Puebla, Oaxaca, Tlaxcala, and Veracruz. Psittaciforrnes. Psittacidae: Rhynchopsitta Synonyms. Serranías Meridionales province p achyrrhynch a (AR). Trogoniformes. (Rzedowski, 1978: 103), Central Mexican oak Trogonidae: Euptil otis neoxenus (AR). forests ecoregion (Dinerstein et al., 1995: 102), Mammalia. Rodentia. Muridae: Microtus Mexican Transvolcanic pine and oak forests mexicanus madrensis, Peromyscus aztecus ecoregion (Dinerstein el al., 1995: 102), Pue- spicilegus (AR, SU); Sciuridae: Glaucomys bla xeric shrublands ecoregion (Dinerstein el volans madrensis, Sciurus nayaritensis apa- al., 1995: 1 10), Transversal Volcanic Axis cheCAR). province (Arriaga el al., 1997: 64; Morrone el al., 1999: 51 1), Oaxaca province (Arriaga et Sierra Madre Oriental province (Fig. 1.2) al., 1997: 65; Morrone el al., 1999: 511), Faja Eastern Mexico, in the states ofSan Luis Po- Volcánica province (Escalante el al., 1998: tosí, Coahuila, Hidalgo, Nuevo León, and 285), and Eje Ne o lvo lcán ico province Querétaro, aboye 1,500 m altitude. (Morrone et al., 1999: 511). Synonyms. Sierra Madre Oriental province Taxa. MAGNOLlOPHYTA. Magnoliop-sida. (Rzedowski, 1978: 103; Arriagaet al., 1997: 64; Asterales. Asteraccae: Montanea frutescens Escalante el al., 1998: 285; Campbell, 1999: (FU). ARTHROPODA. Hexapo-da. 114; Morrone et al., 1999: 511) and Sierra Coleoptera: Passalidae: Odont ot aenius Madre Oriental pine and oak forests ecoregion cuspidatus (CS); Scarabaeidae: Colo/a (Dinerstein et al., 1995: 102). globulicornis, Onthophagus hippopotamus Taxa. MAGNOLIOPHYTA. Liliopsida. (LH, MR). Diptera. Ceratopogonidae: Liliales. Agavaceae: Agave tenuifolia (ZA). Culic oides al bomaculata, C. bakeri, C.

48 Juan J. Morrone dampfi (AR); Mydidae: Mydas oaxacensis (Rzedowski, 1978: 103), Sierra Madre del Sur (AR). Hymenoptera. Formicidae: Polybia pine and oak forests ecoregion (Dinerstein et simillina (AR). VERTEBRAT A. Arnphibia. al., 1995: 103), Guerrero cacti shrublands Anura. Ranidae: Rana montezumae (AR). ecoregion (Dinerstein et al., 1995: 111), Sierra Urodela. Phetodontidae: Pseudoeurycea Madre del Sur province (Arriaga et al., 1997: scandens (AR). Aves. Passeriformes. 65; Campbell, 1999: 116; Morrone et al., 1999: Fringillidae: Geothlypis speciosa (AR). 511), Nudo de Zempoaltépetl province Marnrnalia. Didelphimorphia. Didelphidae: (Escalante et al., 1998: 285), Sierra de Marmosa canescens oaxacae (AR). Mihuatlán province (Escalante et al., 1998: Insectivora. Soricidae: Cryptotis goldmani 285), Sierra Madre del Sur-Oaxaca province alticola, Sorex vagrans orizabae (AR). (Escalante et al., 1998: 285), and Sierra Madre Lagomorpha. Leporidae: Sylvilagus del Sur-Guerrero province (Escalante et al., jloridanus aztecus (AR). Rodentia. Muridae: 1998:285). Peromyscus aztecus hylocetes, P. leucotus, Taxa. MAGNOLIOPHYTA. Magnoliop-sida. Reithrodontomys chrysopsis (AR, SU); Asterales. Asteraceae: Montanoa Sciuridae: Spermophilus m. mexicanus (AR). grandiflora, M. mollissima, M. tomentosa subsp. microcephala (FU). Sapindales. Balsas Basin province (Fig. 1.4) Burseraceae: Bursera aloexylon, B. aptera, Central Mexico, in the states of Guerrero, B. submoniliformis (KS). ARTHROPODA. Mexico, Jalisco, Michoacán, Morelos, Oaxaca, Hexapoda. Coleoptera. Passalidae: Petrejoides and Puebla, below 2,000 m altitude. imbellis, P. jalapensis (CE); Scarabaeidae: Synonyrns. Balsas Basin province Cotinis ibarrai, Onthophagus bassarisus (Rzedowski, 1978: 108; Arriaga et al., 1997: (DE, DY, LH); Staphylinidae: Gansiaflavata 62; Escalante et al., 1998: 285; Morrone et al., (AS). Orthoptera. Pyrgomorphidae: 1999: 511), and Balsas dry forests ecoregion Sphenar ium p. purpurascens (KE). (Dinerstein et al., 1995: 99). VERTEBRA TA. Arnphibia. Anura. Ranidae: Taxa. MAGNOLIOPHYT A. Magnoliop-sida. Rana sierramadrensis (AR). Squarnata. Asterales. Asteraceae: Montanoa liebmannii, Viperidae: Porthidium barbouri (CM). Aves. M. reveali (FU). Fabales. Fabaceae: Trochiliformes. Trochilidae: Amazilia wagleri, Brongniartia montalvoana (DA). Sapindales. Calothorax lucifer, Cynanthus sordidus, Burseraceae: Bursera trimera (AR, RZ). Eupherusa cyanophrys (AR). Marnrnalia. ARTHROPODA. Hexapoda. Coleoptera. Rodentia. Muridae: Peromyscus aztecus Scarabaeidae: Cotinis pueblensis (DE). evides (SU). VE:RTEBRATA. Aves. Passeriformes. Certhiidae: Campylorhynchusjocosus (AR); Mexican Gulfprovince (Fig. 1.6) Fringillidae: Aimophila humeralis (AR). Coast ofthe Mexican Gulf, in eastern Mexico, Strigiformes. Strigidae: Otus seductus (AR). Belize, and northern Guatemala. Marnrnalia. Rodentia. Geomyidae: Ortho- Synonyrns. Lempira- Tegucigalpan province geomys grandis alleni, o. g. carbo, o. g. (Ryan, 1963: 25), Mexican Xerophyllous guerrerensis, o. g. soconuscensis (AR). province (Cabrera & Willink, 1973: 34), Cen- tral American rainforest center (Müller, 1973: Sierra Madre del Sur province (Fig. 1.5) 10), Mexican Gulfprovince (Rzedowski, 1978: South central Mexico, from southern 109; Arriaga et al., 1997: 63; Morrone et al., Michoacán to Guerrero and Oaxaca, and part 1999: 511), Tehuantepec Isthmus moist forests ofPuebla, aboye 1,000 m altitude. ecoregion (Dinerstein et al., 1995: 91), Flooded Synonyrns. Serranías Meridionales province forests ofBelize ecoregion (Dinerstein et al.,

49 Toward a cladistic model for the Caribbean subrcgion

1995: 92), Tamaulipasand Veracruzdry forests 116), and Petén province (Escalante el al., ecoregion (Dinerstein el al., 1995: 99), Veracruz 1998:285). dry forests ecoregion (Dinerstein el al., 1995: Taxa. MAGNOLIOPHYT A. Magnoliop-sida. 99), Veracruz oak forests ecoregion (Dinerstein Sapindales. Burseraceae: Bursera graveolens, el al., 1995: 102), Belize pine and oak forests B. simaruba (AR, KS). Scrophulariales. ecoregion (Dinerstein el al., 1995: 103), Scrophulariaceae: Tetranema roseum (MV). Eastern Mexican grasslands ecoregion ARTHROPODA. Hexapoda. Coleoptera. (Dinerstein el al., 1995: 104), Tabasco and Attelabidae: Pilolab us purpureus, P. Veracruz savannas ecoregion (Dinerstein el splendens (HM); Curculionidae: Caecossonus al., 1995: 104), Veracruz palm savannas sylvaticus (HO); Heteroceridae: Heterocerus ecoregion (Dinerstein el al., 1995: 106), Sierra cr ossi (MI); Pselaphidae: Eutr ich.ites Norte de Chiapas province (Escalante el al., veracruzensis (CF); Scarabaeidae: Cotinis 1998: 285), Sierra de los Tuxtlas province punctatostriata (DE). Lepidoptera. (Escalante el al., 1998: 285; Campbell, 1999: Papilionidae: Troilides tolus (LL); Pieridae:

Fig. 1. Mesoamerican biogeographic provinces ofthe Caribbean subregion. 1, Sierra Madre Occidental; 2, Sierra Madre Oriental; 3, Transmexican Volcanic Belt; 4, Balsas Basin; 5, Sierra Madre del Sur; 6, Mexican Gulf; 7, Yucatán peninsula; 8, Mexican Pacific Coast; 9, Chiapas; 10, Eastern Central America; 11, Western Panamanian Isthmus.

50 Juan l. Morrone Dismorphia eunoe popoluca (LL). Orthoptera. jamaicensis gaumeri (AR). Galliformes. Pyrgomorph idae: Sphenarium m. mexicanum Phasianidae: Colinus nigrogularis, (KE). VERTEBRA TA. Actinopterygii. Meleagris ocellata (AR). Passeriformes. Cyprinodontiformes. Poecilidae: Gambusia Certhiidae: Campylorhynchus yucatanicus eurystoma (LD). Amphibia. Anura. Hylidae: (MU); Tyrannidae: Myarchus yucatanensis Hyla ebraccata (MU). Squamata. Anguidae: (MU). Psittaciformes. Psittacidae: Amazona Celestus eneagr am us (AR); Elapidae: xanthoclora (AR). Strigiformes. Caprimul- Micrurus elegans and M limbatus (CV). Aves. gidae: Caprimulgus badius (AR). Mammalia. Strigiformes. Caprimulgidae: Caprimulgus Carnivora. Procyonidae: Nasua nasua maculicaudus (AR). Mammalia. Didelphi- yucatanica (AR). Primates. Cebidae: Alouatta morphia. Didelphidae: Caluromys derbianus pigra (EM). Rodentia. Muridae: Otonyctomys aztecus (AR). Rodentia. Dasyproctidae: hatti (AA). Dasyprocta mexicana (MU); Muridae: Peromyscus leucopus incensus (AR). Mexican Pacific Coast province (Fig. 1.8) Western Mexico, in the Pacific coast of the Yucatán peninsula province (Fig. 1.7) states of Sinaloa, Nayarit, Colima, Jalisco, Yucatán península, in the Mexican states of Michoacán, Guerrero, Oaxaca, and Chiapas. Campeche, Yucatán, and Quintana Roo, below Synonyms. Escuintla-Usulutan province 200 m altitude. (Ryan, 1963: 24), Chinandegan province (Ryan, Synonyms. Yucatán peninsula province 1963: 27), Mexican Xerophyllous province (Ryan, 1963: 22; Rzedowski, 1978: 109), (Cabrera & Willink, 1973:34), Central American Yucatán center (Müller, 1973: 16),Yucatán dry Pacific center (Müller, 1973: 19), Pacific Coast forests ecoregion (Dinerstein el al., 1995: 99), province (Rzedowski, 1978: 107; Arriaga et al., Yucatán moist forests ecoregion (Dinerstein 1997: 62; Morrone et al., 1999: SI 1), el al., 1995: 91), Quintana Roo wetlands Revillagigedo Islands province (Rzedowski, ecoregion (Dinerstein el al., 1995: 106), 1978: 108; Escalante et al., 1998: 285), Yucatán province (Arriaga el al., 1997: 63; Sinaloense province (Álvarezet al., 1995: 79), Escalante et al., 1998: 285; Morrone el al., Oaxaca moist forests ecoregion (Dinerstein et 1999: 511), Petén province (Arriagaet al., 1997: al., 1995: 91), Sinaloa dry forests ecoregion 63; Morrone et al., 1999: 511), and Cozumel- (Dinerstein et al., 1995: 99), Jalisco dry forests Isla Mujeres province (Escalante el al., 1998: ecoregion (Dinerstein et al., 1995: 99), Pacific 285). Central American dry forests ecoregion Taxa. MAGNOLlOPHYTA. Liliopsida. (Dinerstein et al., 1995: 99), Jalisco palm Poales. Poaceae: Gouninia latifolia var. savannas ecoregion (Dinerstein et al., 1995: guatem alensis, G. p apillosa (OR). 106), Tierras Bajas del Istmo province Magnoliopsida. Fabales. Fabaceae: (Escalante et al., 1998: 285), Southern Costa Caesalp inia gaumer i (AR, RZ). Este province (Escalante et al., 1998: 285), Tres ARTHROPODA. Hexapoda. Coleoptera. Marías Islands province (Escalante el al., Carabidae: Pasimachus purpuratus (NI); 1998: 285), and Pacific Lowlands province Curculionidae: Caecossonus continuus (HO). (Campbell, 1999: 115). Lepidoptera. Papilionidae: Priamides rogeri Taxa. MAGNOLlOPHYTA. Liliopsida. (LL). VERTEBRATA. Squamata. Iguanidae: Poales. Poaceae: Gouninia isabelensis (OR). Enyaliosaurus defensor, Scelop orus Magnoliopsida. Asterales. Asteraceae: cozumelae (AR, SA); Viperidae: Porthidium Montanoa andersonii, M. laskowskii, M. yucatannicum (CV, MU). Aves. standleyi (FU). Sapindales. Burseraceae: Columbiformes. Columbidae: Leptotila Bursera arborea, B. attenuata, B. excelsa (KS,

51 Toward a c\adistic rnodcl for the Caribbean subrcgion

RZ). ARTHROPODA. Hexapoda. Coleop-tera: pine and oak forests ecoregion (Dinerstein et Attelabidae: Euscelus rufiventris (HT); al., 1995: 103), Teh uantepec savannas Curculionidae: Tyloderma affine (WR); ecoregion (Dinerstein et al., 1995: 104), Passalidae: Petrejoides olmecae (CE); Motagua Valley xeric shrublands ecoregion Scarabaeidae: Onthophagus solisi, (Dinerstein et al., 1995: 111), Altos de Chiapas Viridimicus cyanochlorus (GH, lA). Diptera. province (Arriaga et al., 1997: 66), and Chiapas Ropalo-rneridae: Mexicoa mexicana (RH); province (Morrone et al., 1999: 511). Simuliidae: Simulium pseudocallidum, S. Taxa. MAGNOLIOPHYT A. Magnoliop-sida. veracruzanum (CB). Hymenoptera. Apidae: Asterales. Asteraceae: Chionolaena sartorii, Geotrigona acapulconis (CD). Lepidoptera. Montanoa echinacea, M. guatemalensis, M. Papilionidae: Battus eracon, ilus pteropoda (FE, FU). Haloragales. occiduus, Pyrrhostica abderus baroni (LL); Gunneraceae: Gunnera kilipiana (RZ). Pieridae: Dismorphia amphiona lupita, D. a. Scrophulariales. Scrophulariaceae: Tetranema isolda, D. crisia alvarezi, Enantia fina ssp., evolutum (MV). Urticales. Cecropiaceae: Pieriballia viardi, Prestonia clarki (LL). Cecropia sylvicola (FR). ARTHROPODA. Orthoptera. Pyrgomorphidae: lchthiacris Hexapoda. Coleoptera: Cerambycidae: Anejlus elongata(KE). VERTEBRATA. Amphibia. poriferus, Eburia schusteri, Megapsyrassa Anura. Hylidae: Hyla robertmertensi, H. testacea, M. chiapaneca, Oxycoleus piceus, sartori (MU, SA). Squamata. Viperidae: Pachymerola wappesi, Rhodoleptus Bothrops langsbergi aphryomegas, Crotalus nigripennis, Semanotus australis (G B); basiliscus (CV, MU). Aves. Craciforrnes. Cleridae: Enoclerus gabriellae (RI); Cracidae: Ortalis wagleri (AR). Galliformes. Passalidae: Ogypes, Petrejoides guatemalae Phasianidae: Callipepla doug/asi (AR). (CE, SR); Scarabaeidae: Ap elt astes Psittaciformes. Psittacidae: Amazona finschi ehiapasensis, Geotrupes pilanolensis, Golofa (AR). Mammalia. Lagornorpha, Leporidae: ch am p io ni, Hologymnetis kinichahau, Lepus flavigularis (MU). Rodentia. Trigonop/etastes glabella, Viridimicus Geomyidae: Orthogeomys grandis sspp. (AR); aurescens (HD, iu, HK, lA, MR, RO); Sciuridae: Sciurus colliaei (AR). Staphylinidae: Gansia andersoni, G. fortemaculata, G. tibialis, Styngetus Chiapas province (Fig. 1.9) championi (AS, NH). Heteroptera. Miridae: Mexico, Guatemala, and Nicaragua, basically A tractotomus teopisca (SS). Lepidoptera. corresponding to th e Sierra Madre de Papilionidae: Baronia brevicornis Chiapas, from 500 to 2,000 m altitude. rufodiscalis, Parides panares lycimenes, Synonyms. Chiapan-Guatemalan Highland Priamides e. erostratus (LL); Pieridae: Colias province(Ryan, 1963: 23; Campbell, 1999: 116), philodice, Dismorphia eunoe chamula, D. e. Nicaraguan Montane province (Ryan, 1963: eunoe, Perrhybris pamela chajuelnsis, 28), Central American montane forest center Pseudopieris nehemia irma (LL). Orthoptera. (Müller, 1973: 14), Guatemalan montane forest Pyrgomorphidae: Prosphena scudderi, subcenter (Mü ller , 1973: 14), Serranías Sphenarium mexicanum histr io (KE). Transístmicas province (Rzedowski, 1978: VERTEBRA TA. Amphibia. Anura. Ranidae: 103), Soconusco province (Rzedowski, 1978: Rana maculata (AR). Urodela. 109; Arriaga et al., 1997: 66; Morrone et al., Plethodontidae: Nototriton b arbouri, N. 1999: 511), Sierra Madre moist forests nasalis, N. veraepacis (PF). Squamata. ecoregion (Dinerstein et al., 1995: 92), Central Iguanidae: Enyaliosaurus palearis, American montane forests ecoregion Sceloporus malachitichus (SA); Anguidae: (Dinerstein et al., 1995: 92), Central American Abronia ochoterenai (AR). Aves.

52 Juan J. Morrone

Passeriformes. Furnariidae: Dendrocolaptes lia. Rodentia. Muridae: Oryzomys devius (MU). picumnus (AR); Muscicapidae: Turdus plebejus (AR). Strigiformes. Strigidae: Otus Western Panamanian Isthmus province (Fig. barb atus (AR). Mammalia. Rodentia. l.ll) Muridae: Peromyseus aztecus oaxacensis Western Central America, from Costa Rica to (SU). western Pan ama. Synonyms. Puntarenas-Chiriquí province Eastern Central America province (Fig. 1.10) (Ryan, 1963: 30), Chiriquí subcenter (Müller, Eastern Central América, from Guatemala to 1973: 24), Pacific Panama Isthmus moist forests Panama. ecoregion (Dinerstein et al., 1995: 92), Panama Synonyms. Mosquito province (Ryan, 1963: dry forests ecoregion (Dinerstein et al., 1995: 29), Guatuso- Talamancan province (Ryan, 100), and Costa Rican-western Panamanian 1963: 31), Talamanca montane forest subcenter province (Campbell, 1999: 116). (Müller, 1973: 14), Coco center(Müller, 1973: Taxa. ARTHROPODA. Hexapoda. Coleop- 22), Costa Rican center (Müller, 1973: 23), tera. Passalidae: Petrejoides tenuis (CE); Mosquito subcenter (Mü ller , 1973: 24), Scarabaeidae: Hemiphileurus jamesoni, H. Talamanca paramo center (Müller, 1973: 26), youngi, Onthophagus orphnoides (GH, RL); Atlantic Central American moist forests Staphylinidae: Gansia obscura, G. ecoregion (Dinerstein et al., 1995: 92), tergopunctata,G. unizonata(AS); Tenebrionidae: Talamanca montane forests ecoregion Archaeoglenes bollensis, A. puntaensis (WA). (Dinerstein et al., 1995: 92), Moisquitía pine Diptera. Simuliidae: Simulium panamense (CB). forests ecoregion (Dinerstein et al., 1995: 103), Costa Rica Paramo ecoregion (Dinerstein et Bahama province (Fig. 2.1) al., 1995: 107), and Eastern Panamanian Bahama archipelago, comprising the islands Highlands province (Campbell, 1999: 116). of Abaco-Grand Bahama, Andros-Bimini, Cat, Taxa. MAGNOLlOPHYT A. Magnoliop-sida. Crooked-Mayaguana, Exumas, Inaguas, . : sect. Long-Ragged Island Range, Mona, New Jimenezia (BR). ARTHROPODA. Hexapo-da. Providence-Eleutheras, San Salvador-Rum Coleoptera. Curculionidae: Hammatos-tylus Cay, St. Eustatius, St. Kitts, St. Lucia, St. cr iniger, H. exiguus, Sicoderus Martin, St. Vincent, and Turks and Caicos. appendiculatus, Tyloderma expansum, T. Synonyms. Caribbean province (Cabrera & lepidogram ma (VN, WR); Passalidae: Willink, 1973: 38), Bahamas dry forests Petrejoides subrecticornis (CE); Pselaphidae: ecoregion (Dinerstein et al., 1995: 98), and Eutrichites buscki, E. funiculis (CF); Bahamas pine forests ecoregion (Dinerstein Staphylinidae: Gansia taeniata (AS); etal., 1995: 102). Tenebrionidae: Archaeoglenes occidentalis Taxa. ARTHROPODA. Hexapoda. Coleop- (W A). Hymenoptera. Apidae: Geotrigona tera. Carabidae: Glyptolenus smithi (LO); chiriquiensis (CD). VERTEBRAT A. Acti- Curculionidae: Decuanellus bahamensis nopterygii. Cyprinodontiformes. Poecilidae: (HO). Diptera. Drosophilidae: Drosophila Gambusia nicaraguensis (LO). Amphibia. antillea, D. insularis, D. vittatifrons, Stegana Anura. Hylidae: Hyla phlebodes, H. urano- horae, S tarsalis (GI). Hemiptera. Lygaeidae: chroa (MU). Urodela. Plethodontidae: Baihydema socia, Ochrimnus laevus, Ozophora Nototri-ton picadoi, N. richardi (PF). octomaculata (SL). VERTEBRATA. Squamata. Viperidae: Bothrops langsbergi Actinopterygii. Cypri no-donti form es. annectens (MU); Elapidae: Micrurus alleni, M Cyprinodontidae: Cyprinodon laciniatus multifasciatus (CV); Viperidae: Bothriechis (RA, RV); Poecilidae: Gambusia hubbsi, G. /ateralis, Porthidium picadoi (CV). Mamma- 53 Toward a c\adistic model for the Caribbcan subrcgion manni (LO). Ophidiiformes. Bythitidae: Polycentropidae: Polycentropus nigriceps Lucifuga spelaeotes (RV). Chelonia. (HL). VERTEBRATA. Actinopterygii. Emydidae: Trachemys stejnegeri malonei Cyprino-dontiformes. Cyprinodontidae: (SE). Squamata. Boidae: Epicrates Cubanichthys cubensis, Fundulus grandis, chrysogaster, E. exsul, E. monensis (KL, PO); Girar dinus, Quintana, Rivulus Colubridae: Chironius vincenti, Clelia insulaepinorum, R. garciai (RA, RV); errabunda, Mastigodryas bruesi (HB); Poecilidae: Gambusia pune/ata, G. Leptotyphlopidae: Leptotyphlops columbi puncticulata, Poecilia vittata (LO, RA, RV). (HB); Teiidae: Cnemidophorus vanzoi (HB); Lepisosteiformes. Lepisosteidae: Atractosteus Viperidae: Bothrops caribbaeus, B. tristoechus (RA, RV). Perciformes. Cichlidae: lanceolatus (CV). Cichlasoma ramdensi, C. tetracanthus (RA). Synbranchiformes. Synbranchidae:Ophisternon Cuba province (Fig. 2.2) aenigmaticum (RV). Chelonia. Emydidae: Island ofCuba. Trachemys d. decussata, T d angusta (SE). Synonyms. Caribbean province (Cabrera & Squamata. Boidae: Epicrates angulifer (KL, Willink, 1973: 38), Cuban moist forests PO); Gekkonidae: Sphaerodactylus spp., ecoregion (Oinerstein et al., 1995: 90), Cuban Tarentola americana (HB, PG, PO); dry forests ecoregion (Oinerstein el al., 1995: Iguanidae: Chamaeleolis (GY); Xantusiidae: 98), Cuban pine forests ecoregion (Oinerstein Cricosaura (CT). Crocodylia. Crocodylidae: el al., 1995: 101), Cuban wetlands ecoregion Crocodylus rhombifer (HB). (Oinerstein et al., 1995: 105), and Cuban cacti shrublands ecoregion (Oinerstein et al., 1995: Cayman islands province (Fig. 2.3) 109). Archipe1ago including Grand Cayrnan, Little Taxa. ANNELIDA. Oligochaeta. Megas- Cayman, and Cayman Brac. colecidae: Zapatadrilus morenoae, Z. siboney, Synonyms. Caribbean province (Cabrera & Z. taina (RF). ARTHROPODA. Hexapoda. Willink, 1973: 38), Cayman islands dry forests Blattaria. Blaberidae: Epilampra spp. (GU); ecoregion (Oinerstein el al., 1995: 98), and Blattellidae: Nesomylacris spp. (GU); Blatidae: Cayman islands xeric shrublands ecoregion Eurycotis spp. (GU). Coleoptera. Carabidae: (Oinerstein et al., 1995: 109). Clivina cubae, Scarites alternans, S. cubanus Taxa.ARTHROPODA. Hexapoda. Hemiptera. (NI); Curculionidae: Caecossonus decuanus, Lygaeidae: Ochrimnus nigriceps, Ozophora Decuanellus vinai, Neomastix ver itas, miniscula, o. pallidifemur (SL). VERTE- Pseudocaec ossonus zayasi, BRAT A. Actinopterygii. Cyprinodon- Pseudo alaocyb ites negreai, Sicoderus tiformes. Po ec ili dae: Gamb usia sleeper i (CR, HO, VN); Latridiidae: caymancensis, Limia caymanensis (LO, RV). Metophthalmus cuba (AP); Scarabaeidae: Hemiphileurus cribratus, H cubaensis (RE); Jamaica province (Fig. 2.4) Sphindidae: Carinisphindus bicolor (MC). Is1and of Jamaica. Oiptera. Tephritidae: Anastrephas insulae, A. Synonyms. Caribbean province (Cabrera & soroana (FN). Hemiptera. Cicadidae: Juanaria Willink, 1973: 38), Jamaican moist forests (RM); Lygaeidae: Kleidoceris suffusus, ecoregion (Oinerstein el al., 1995: 91), and Lygaeus dearmasi, L. wygodzinskyi, Jamaican dry forests ecoregion (Oinerstein el Melanopleurus tetr asp ilus, Patritius al., 1995: 98). cubensis (SL); Membracidae: Stalotypa (RM). Taxa. ARTHROPODA. Hexapoda. Coleop- Hymenoptera. Formicidae: Codioxenus tera. Buprestidae: Agrilus [amaicensis (HS); simulans, Dorisidris nitens (WI). Trichoptera. Carabidae: Ardistomis franki, Glyptlenus

54 .Juan .J. Morrone latelytra, Platynus cinchonae, P. faber, P. Synonyms. Caribbean province (Cabrera & jamaicae, P. rastafarius species groups (LE, Willink, 1973: 38), Hispaniola moist forests L1, LO, NI); Coccinellidae: Psorolymasicardi ecoregion (Dinerstein et al., 1995: 90), (GR); Curculionidae: Pseudoalaocyobites Hispaniola dry forests ecoregion (Dinerstein affinis, P. armatus, P. diversesculptus, P. el al., 1995: 98), Hispaniola pine forests inermis, Pcjarmilae, P. pacei, P. persimilis, P. ecoregion (Dinerstein et al., 1995: 102), and stewartii (HO); Scarabaeidae: Hemiphileurus Lago Enriquillo Cuban wetlands ecoregion jamaicensis (RE); Sphindidae: Carinisphindus (Dinerstein et al., 1995: 106). platysphinctos (MC); Tenebrionidae: Taxa. MAGNOLlOPHYT A. Magnoliop-sida. Archaeoglenes pecki (WA). Diptera. Myrtales. Onagraceae: Fuchsia triphylla, F. Drosophilidae: Chymomyza jamaicensís, pringsheimii (BR). ARTHROPODA. Drosophila paraguttata (G 1). Hemiptera. Chilopoda. Geophilomorpha: Schendylidae: Lygaeidae: Oncope/tus spectabilis (SL); Ctenophilus nes iotes (PD). Hexapoda. Membracidae: Quadrinareini (RM). Coleoptera: Anthribidae: Phaenotheriopsis Hymenoptera. Formicidae: Strumigenys tuberculatus, P. umbonatus, P. verrucosus jamaicensis (LG). Trichoptera. Polycentropidae: (VA); Carabidae: Antilliscaris darlingtoni, Po/ycentropus jamaicensis (HL). VERTE- Barylaus puncticeps, P/atynus biramosus, P. BRA TA. Actinopterygii. Cyprinodontiforrnes. cristophe, P. jaeger i, P. laeviceps, P. Poecilidae: Gamb usia melapleura, G. transcibao species groups (LA, LE, L1, NI); oligosticta, G. wrayi (LD, RV). Chelonia. Coccinellidae: Bura, Psorolyma doyeni, P. Emydidae: Trachemys terrape n (SE). baorucensis, P. cyanella (GR); Curculionidae: Squamata. Boidae: Epicrates subflavus (KL, Kuschelaxius disc ifer, Micromyrmex PO); Gekkonidae: Sphaerodactylus parkeri, asclepia, Sicoderus championi, S. ramosi, S. S. richardsoni, S. semasiops (PG, PO). truncatipennis (HO, IS, VN); Lathridiidae: Metophthalmus columbusi, M schusteri, M Hispaniola province (Fig.2.5) trilineatus (AN); Micropeplidae: Pep/omicrus Island ofHispaniola (Dominican Republic and iviei (CN); Passandridae: Cat ogenus Haiti). slipinskii (TH); Scarabaeidae: Hemiphileurus

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7 -

11

Fig. 2. Antillean biogeographic provinces ofthe Caribbean subregion. 1, Bahama; 2, Cuba; 3, Cayman islands; 4, Jamaica; 5, Hispaniola; 6, Puerto Rico; 7, Lesser Antilles. 55 Toward a c1adistic rnodel for (he Caribbean subrcgion dispar, H. laeviceps, H. phratrius, H. ryani, Hemiphileurus puertoricensis (RE). Diptera. H. scutellaris (RE); Sphindidae: Drosophilidae: Mayagueza argentifera (G 1). Car inisphindus leptos-phinctos (MC). Hemiptera. Cicadidae: Borencona (RM); Diptera. Drosophilidae: Drosophila nesiota Lygaeidae: Ochrimnus henryi (SL); (GI). Embioptera. Teratembiidae: Oligembia Membracidae: Jibarita (RM). Trichoptera. vetusta (SZ). Hemiptera. Lygaeidae: Polycentropidae: Polycentropus zaneta (HL). Melanopleurus maculicorium, Pamphantus VERTEBRAT A. Chelonia. Emydidae: spp. (SL); Tibicinidae: Psallodia (RM). Trachemys s. stejnegeri (SE). Squamata. Hymenoptera. Formicidae: Hypocryptocerus Boidae: Epicrates inornatus (KL, PO). haemorrhoidalis (WI). Orthoptera. Acrididae: Amblytropidia (PL). Trichoptera. Lesser Antilles province (Fig. 2.7) Polycentropidae: Polycen-tropus criollo, P. Archipelagos ofthe Lesser Antilles (Antigua, domingensis, P. marcanoi(HL). VERTEBRATA. Barbados, Barbuda, Desirade, Dominica, Actinopterygii. Cyprinodon-tiformes. Grenada, Guadeloupe, Marie Galante, Cyprinodontidae: Cyprinodon bondi (RA, RV); Martinique, Montserrat, Nevis, and Saba) and Poecilidae: Curtipenis, Limia, Poecilia the Virgin islands (Anegada, Culebra, St. Croix, (Odontolimia), P. dominicensis, P. elegans, P. St. John, St. Thomas, Tortola, Vieques, and hispaniolana(RA, RV). Perciforrnes. Cichlidae: Virgin Gorda). Cichlasoma haitiensis (RA). Amphibia. Anura. Synonyms. Caribbean province (Cabrera & Leptodactyl idae: Eleutherodactylus Willink, 1973: 38), Barlovento islands moist (Pelorius) (LC). Chelonia. Emydidae: forests ecoregion (Dinerstein et al., 1995: 91), Trachemys decorata, T stejnegeri vicina (SE). Sotavento islands moist forests ecoregion Squamata. Boidae: Epicrates fordii, E. (Dinerstein et al., 1995: 91), Barlovento islands gracilis (KL, PO); Gekkonidae: dry forests ecoregion (Dinerstein et al., 1995: Sphaerodactylus spp. (PG); Iguanidae: 99), Sotavento islands dry forests ecoregion Chamaelinorops (GY); Scindidae: Mabuya (Dinerstein et al., 1995: 99), Barlovento islands lineolata (HB). xeric shrublands ecoregion (Dinerstein et al., 1995: 109), and Sotavento islands xeric Puerto Rico province (Fig.2.6) shrublands ecoregion (Dinerstein et al., 1995: Island ofPuerto Rico. 109). Synonyms. Caribbean province (Cabrera & Taxa. ARTHROPODA. Chilopoda. Willink, 1973: 38), Puerto Rico moist forests Geophilomorpha. Geophilidae: Ityphilus ecoregion (Dinerstein et al., 1995: 91), and mauriesi, Taeniolinum setosum guade- Puerto Rico dry forests ecoregion (Dinerstein loupensis (DP). Hexapoda. Coleoptera: et al., 1995: 98). Carabidae: Ardistomis atr ip ennis, A. Taxa. ARTHROPODA. Hexapoda. Coleop- guadeloupensis, Clivina aff. latiuscula, C. tera: Anthribidae: Ph aenother iop sis tuberculata, Glyptolenus simplicicollis, conciliatus (V A); Carabidae: Antilliscaris Platynus alternans, P. ellipticus, Stratiotes danforthi, A. megacephalus, A. mutchleri, anglicanus, S. iracundus (LE, LI, LO, NI); Barylaus estriatus, Oxydrepanus coamensis Curculionidae: Decuanellus brevicrus, D. (LA, NI); Coccinellidae: Lenasa jayuyai, buclavatus, D gladiatus, D. muchmorei, D. Neaptera doyeni, N. viridissima, Psorolyma viti, Sicoderus contiguus, S. delauneyi, S. maxillosa (GR, GS); Curculionidae: Decua- remotus (HO, VN); Lathridiidae: Methophtha nellus longirostris, D. pecki, Kuschelaxius musiviei, M. muchmorei (AP). Diptera. tomentosus (HO); Lathridiidae: Metoph- Drosophilidae: Drosophila hypophallus, D. thalmus rectangulatus (AN); Scarabaeidae: insularis, D. nigrodunni (GI). Hemiptera.

56 Juan J. Morronc Lygaeidae: Blissusplanus (SL). Trichoptera. Rourea antioquensis (FO). Sapindales. Polycentropidae: Polycentropus insularis Zygophyllaceae: Bulnesia arborea (HH). (HL). VERTEBRATA. Amphibia. Dendro- Violales. Passifloraceae: Passiflora bracteosa, batidae: Colostethus chalcopis (HB). P. purdiei, P. rugosa var. rugosa, P. Squamata. Colubridae: Liophis cursor species schlim.iana (ES). ARTHROPODA. group (HB); Gekkonidae: Phyllodactylus Arachnida. Araneae. Clubionidae: pulcher, Sphaerodactylus fantasticus, S. Trachelopachys magdalena (PB). Opiliones. sab anus, S. sputator (HB, PG, PO); Stygnidae: Eutimesius albicinctus, Innoxius Gymnophthalm idae: Gymnophthalmus pleei magnus, Stygnus aggerum (PI). Crustacea. (HB); Iguanidae: Iguana delicatissima (HB). Decapoda. Pseudotelphusidae: Chaceus davidi, C. nasutus, Orthothelphusa holthuisi, Maracaibo province (Fig. 3.1) Strengeriana taironae (CH, RC); Northern Colombia and northwestern Vene- Trichodactylidae: Bottiella niceforei (MG, zuela. ML). Hexapoda. Coleoptera. Curculionidae: Synonyms. Barranquilla centre (Müller, 1973: Ludovix bifasciatus, Macrostylus beatricis, 28), Santa Marta centre (Müller, 1973: 30), M. lacunitae, M. mucuyensis, M. valeranus, Catatumbo centre (Müller, 1973: 52), Naupactus rosalesi, Sicoderus hirsutus (BD, Maracaibo subcentre (Müller, 1973: 57), Sie- BO, VN); Staphylinidae: Neobisniusfortis and ITaNevada centre (Müller, 1973: 31), Maracaibo N. vigii (FA); Trogidae: Omorgus badeni (SC). province (Ringuelet, 1975: 107), Nechí area VERTEBRAT A. Actinopterygii. Chara- (Cracraft, 1988: 223), Catatumbo refuge ciformes. Characidae: Creagrutus (Hernández et al., 1992c: 95), Alta Guajira hildebrandi, C. mar acaib oensis, C. district (Hernández el al., 1992c: 115), Sierra nigr ostigm atus, C. para/acus (HV); Nevada de Santa Marta province (Hernández Curimatidae: Cur imata miv art ii, et al., I992a: 117), Catatumbo moist forests Cyphocharax aspilos (VI, VT). Amphibia. ecoregion (Dinerstein el al., 1995: 97), Santa Anura. Leptodactylidae: Eleuthe-rodactylus Marta montan e forests ecoregion (Dinerstein carmelitae, E. delicatus, E. insignitus, E. el al., 1995: 97), Sinú valley dry forests megalops, E. sanctamartae (MU); Pipidae: ecoregion (Dinerstein et al., 1995: 100), Pipa parva (BZ). Squamata. Elapidae: Maracaibo dry forests ecoregion (Dinerstein Micrurus dissoleucus nigrirostris (MU); et al., 1995: 101),Guajira/Barranquillaxeric Viperidae: Bothrops l. lansbergii (MU). Aves. ecoregion (Dinerstein el al., 1995: 111), Columbiformes. Columbidae: Columba Sierra Nevada de Santa Marta (Rangel el al., corensis (HG). Coraciformes. Momotidae: 1995b: 107), Atlantic coast subregion (Rangel Momotus momota spatha (HG). Craciformes. el al., 1995c: 21), Guaj ira subregion (Rangel el Cracidae: Crax alberti (MU). Passeriformes. al., 1995c: 21), Santa Marta province (Morrone, Certhiidae: Campylorhynchus griseus (MU); 1999:4), Barranquilla province (Morrone, 1999: Formicariidae: Grallaria bangsi (MU); 4), and Maracaibo province (Morrone, 1999: Fringillidae: Anisognathus melanogenys, 4). Atlapetes melanocephalus, Basileuterus Taxa. MAGNOLlOPHYT A. Magnoliop-sida. basilicus, Catamenia homochroa oreophila, Asterales. Asteraceae: Chionolaena Conirostrum rufum, Myioborus .flavivertex chrysocoma, C. columbiana, Montanoajosei (M U); Furnariidae: Cranioleuca hellmayri, (FE, FU). Fabales. Fabaceae: Crotalaria Synallaxis c. candei, S. fuscorufa (MU); viiellina, C. verrucosa, Prosopis juliflora Muscicapidae: Turdus leucolemas cautor (BE, HH). Rhamnales. Rhamnaceae: Condalia (HG); Tyrannidae: Myiotheretes pernix, henriq uezii (FB). Rosales. Connaraceae: Myiarchus panamensis (MU). Piciformes.

57 Toward a c1adistie modcl for the Caribbean subregion

Rhamphastidae: Rhamphastos citreolaemus Coleoptera. Carabidae: Clivina oblita (NI); (HH). Psittaciformes. Psittacidae: Aratinga p. Curculionidae: Macrostylus coloniae, M. pertinax, Pyrrhura viridicata (MU). loscaroniensis, M. pittieri, M. santanae, Tinamiformes. Tinamidae: Crypturellus Naupactus cupreus, N. penai, N. litoris, N. erythropus idoneus (MU). Trochiliformes. santanae, and N. venezolanus, Pseudoa- Trochilidae: Anthocephala floriceps, laocybites e!egans, P. margheritae; and P. Campylopterus phainopeplus, Coeligena squamirostris, Sicoderus abbreviatus, S. phalerata (MU). Marnrnalia. Lagomorpha. bicolor, S. cracens, S. globulicollis (BD, BO, Leporidae: Sylvilagus brasiliensis sancta- HO, VN); Staphylinidae: Cylindroxystus martae, S. floridanus superciliaris (MU). lyrus, Neolindus brachiatus, N. rudiculus Primates. Callithrichidae: Saguinus o. oedipus (HN). Diptera. Sciaridae: Rhipidita (EM). Rodentia. Echimyidae: Proechimys brevicornis, R. vespertilio (AM); Simuliidae: canicollis, P. mincae (PT); Erethizontidae: Simulium clownsi, S. oviedoi species group Coendou bicolor sanctamartae (MU); (CB, CO). Hemiptera. Miridae: Rhinacloa Heteromyidae: Heteromys anomalus jesupi puertoricensis (SV). VERTEBRATA. (MU); Muridae: Oryzomys albigularis Actinopterygii. Characiformes. Curimatidae: maculiventer (MU); Sciuridae: Sciurus Curimata incompta (VI). Cyprinodontiformes. granatensis gerrardi (MU). Rivulidae: Rachovia (PA). Crocodylia. Crocodylidae: Crocodylus acutus (DO). Aves. Coastal Venezuela province (Fig. 3.2) Psittaciformes. Psittacidae: Amazona Northern Venezuela and Colombia. It also barbadensis (MU). Marnrnalia. Didelphi- includes the islands of Aruba, Curacao, and morphia. Didelphidae: Gracilinanus marica Bonaire. (EM). Rodentia. Echimyidae: Proechimys Synonyrns. Caribbean centre (Müller, 1973: guairae and P. urichi (PT). 57), Venezuelan coastal forest centre (Müller, 1973: 54), Venezuelan montane forest centre Trinidad and Tobago province (Fig. 3.3) (Müller, 1973: 56), Caribbean coast province Islands ofTrinidad and Tobago. (Ringuelet, 1975: 107), Araya and Paria xeric Synonyrns. Trinidad province (Ringuelet, scrub ecoregion (Dinerstein el al., 1995: 111), 1975: 107), Trinidad and Tobago dry forests Cordillera de la Costa montane forests ecoregion (Dinerstein etal., 1995: 100), Trini- ecoregion (Dinerstein et al., 1995: 92), Aruba, dad and Tobago moist forests ecoregion Curacao , and Bonaire cacti shrublands (Dinerstein et al., 1995: 93), and Trinidad and ecoregion (Dinerstein et al., 1995: 111), La Tobago province (Morrone, 1999: 4). Costa xeric shrublands ecoregion (Dinerstein Taxa. ARTHROPODA. Arachnida. Opilio- et al., 1995: 111), Lara/ Falcon dry forests nes. Stygnidae: Stygnoplus clavotibialis (PI). ecoregion (Dinerstein et al., 1995: 101), Hexapoda. Coleoptera. Carabidae: Nyctos- Paraguana restingas ecoregion (Dinerstein el tyles planicollis (NI); Coccinellidae: al., 1995: 112), Paraguana xeric scrub ecoregion Nexophallus popei (GN); Curculionidae: (Dinerstein et al., 1995: 111), and Coastal Sicoderus propinquus, Tyloderma pallidum Venezuelan province (Morrone, 1999: 4). (VN, WR). Diptera. Drosophilidae: Droso- Taxa. MAGNOLlOPHYT A. Magnoliop-sida. phila bedicheki (G 1). Trichoptera. G losso- Asterales. Asteraceae: Montanoa fragans somatidae: Protoptila ignera (FL). (FU). ARTHROPODA. Arachnida. Opilio- VERTEBRAT A. Marnrnalia. Rodentia. nes. Stygnidae: Stenostygnellus macrochelis, Echimyidae: Proechimys trinitatus (PT). Stygnoplus biguttatus, S. granulosus, S. meinerti, S. trilineatus (PI). Hexapoda. Chocó province (Fig. 3.4)

58 J uan J. Morrone

Pacific coast ofnorthern Ecuador, Colombia, VERTEBRA TA. Aetinopterygii. Chara- and Panama. ciformes. Characidae: Creagrutus affinis (HV); Synonyms. Colón-Darién province (Ryan, Curimatidae: Cyphocharax magda-lenae, 1963: 32), Colombian Pacific centre (Müller, Pseudocur imatella lineop unctata, 1973: 38), Northern Pacific province Steindachnerina atratoensis (VR, VS, VT). (Ringuelet, 1975: 107), Chocó are a (Cracraft, Amphibia. Anura. Hylidae: Gastrotheca 1988: 223), Chocó-Magdalena provin- cornut a (DL). Chelonia. Bataguridae: ce (Hernández el al., 1992a: 118), Chocó sec- Rhinoclemmys spp. (BU). Squamata. Elapidae: tor (Hernández el al., 1992a: 118), Colombian Micrurus ancor al is (CV); Viperidae: province (Rivas-Martínez & Navarro, 1994: Bothr iopsis punctat a (CV). Aves. map), Chocó/ Darién moist forests ecoregion Columbiformes. Columbidae: Columba (Dinerstein el al., 1995: 96), Eastern Panama goodsoni, C. subvinacea berlepschi (HG). montano forests ecoregion (Dinerstein el al., Craciformes. Cracidae: Penelope ortoni (HG, 1995: 96), Pacific coast region (Rangel el al., MU). Galbuliformes. Galbulidae: Ga/bula 1995a: 121), Pacific coast region (Rangel et ruficauda melanogenia (HG). Galliformes. al., 1995c: 21), Central subregion (Rangel et Phasianidae: Odonthophorus crythr ops al., 1995c: 21), Northern subregion (Rangel et paramb ae (HG). Gruiformes. Rallidae: al., 1995c: 21), Pacific Basin subregion (Rangel Aram ides wolfi (HG). Passeriformes. et al., 1995c: 21), Western subregion (Rangel Fringillidae: Chrysothlypis chrysomelas, C. el al., 1995c: 21), and Chocó province salmoni (MU); Furnariidae: Cranioleuca (Morrone, 1999: 5). erythrops griseigularis (HG); Taxa. CONIFEROPHYTA. Coniferopsida. Thamnophilidae: Sip ia berlep sch.i, S. Con iferales. Podocarpaceae: Podocarpus ros enbergi (MU); Tyrannidae: gu atemalensis var. allenii (TO). Machaeropterus deliciosus (MU). Picifonnes. MAGNOLIOPHYTA. Liliopsida. Arecales. Picidae: Picumnus olivaceus harteni (HG). Arecaceae: Wettinia p anamensis (BE). Psittaciformes. Psittacidae: Pionopslu.a Cyclanthales. Cyclanthaceae: Dicranopygium pulchra (CP). Trochiliformes. Trochilidae: arusisense, D. gigantea, D. odoratum, D. Androdon aequatorialis, Coeligena wi/soni, tril obulata (TB). Magnoliopsida. Aristo- Hy/ocharis grayi humboldtii, Thaluronia lochiales. Aristolochiaceae: Aristolochia furcata fannyi (HG, MU). Trogoniformes. colossifolia. A. trianae (GO). Fabales. Trogonidae: Trogon massena australis, T. Fabaceae: Crotalaria spectabilis (BE). Halo- comptus (HG). Mammalia. Chiroptera. ragalcs. Gunneraceae: Gunnera atropurpurea Phyllostomatidae: Platyrrhinus chocoensis var. chocoana (MD). Myrtales. Onagraceae: (A V). Didelphimorphia. Didelphidae: Fuchsia putumayensis, F. sessilifolia (BR). Caluromys derb ianus (EM). Primates. Rosales. Connaraceae: Connarus nervatus, Callithrichidae: Saguinus geojfroyi (EM); C. silvanensis, C. williamsi, Pseudoconnarus Cebidae: Alouatta palliata, Ateles fuscipes agelaefolius; Rourea pittieri (FO). (EM). Rodentia. Echimyidae: Proechimys ARTHROPODA. Araehnida. Scorpiones. colombianus (PT); Geomyidae: Orthogeomys Buthidae: Ananteris gorgonae (LF). dariensis, O. thaeleris (HF). Crustacea. Decapoda. Trichodactylidae: Melocarcinus meekei (MT). Hexapoda. Cauea provinee (Fig. 3.5) Colcoptera. Scarabaeidae: Onthophagus Colombia and Ecuador. b arr ett i (GH). Diptera. Ephydridae: Synonyms. Cauca centre (Müller, 1973: 33), Mimapsilopa metatarsata, M. schildi (LZ); Colombian montane forest centre (Müller, Simuliidae: Simulium sanguineum (CC). 1973: 34), North Andean centre (Müller, 1973:

59 Toward a cladistic model for thc Caribbean subregion 45), East Andean subcentre (Müller, 1973: 35), (MU). Lagomorpha. Leporidae: Sy/vilagus West Andean subcentre (Müller, 1973: 35), br as iliensis andinus (MU). Rodentia. Cauca valley dry forests ecoregion (Dinerstein Sciuridae: Sciurus pucheranii (MU). et al., 1995: 100), Cauca valley montane forests ecoregion (Dinerstein el al., 1995: 100), Magdalena province (Fig. 3.6) Northwestern Andean montane forests Western Venezuelaand northeastern Colom- ecoregion (Dinerstein et al., 1995: 97), Patía bia. valley dry forests ecoregion (Dinerstein et al., Synonyms. Magdalena centre (Müller, 1973: 1995: 100), and Cauca province (Morrone, 32), Magdalena domain (Ringuelet, 1975: 107), 1999:5). Northern Andean province (Ringuelet, 1975: Taxa. MAGNOLlOPHYT A. Magnoliop-sida. 107), Chocó-Magdalena provin- Faba1es. Fabaceae: Crotalaria brevidens, C. ce (I-Iernández et al., 1992a: 118), Magdalena juncea, C. paulina (BE). I-Ialoragales. sector (I-Iernández et al., 1992a: 125), Gunneraceae: Gunnera atropurpurea var. Colombian province (Rivas-Martínez & Na- sibundoya, G. cuatrecasii, G. manicata, G. varro, 1994: map), Cordillera Oriental montane tamarensis (MD). Malvales. Bombacaceae: forests ecoregion (Dinerstein et al., 1995: 97), Matisia longiflor a (FC). Myrtales. Magdalena/ Uraba moist forests ecoregion Onagraceae: Fuchsia dependens and F. (Dinerstein et al., 1995: 97), Magdalena valley loxenis species groups, F. palIescens, F. dry forests ecoregion (Dinerstein et al., 1995: orientalis (BR). Urticales. Cecropiaceae: 100), Magdalena valley montane forests Cecropia megastachya (FR). Violales. ecoregion (Dinerstein et al., 1995: 100), Mag- Passifloraceae: Passiflora antioquensis, P. dalena province (Morrone, 1999: 4), and Mag- fimbriatistipula, P.flexipes, P. parritae (ES). dalena valley province (Morrone, 1999: 5). ARTHROPODA. Arachnida. Araneae. Taxa. FILlCOPHYTA. Filicopsida. Filica-Ies. Oxyopidae: Peucetia cauca (LR); Opiliones. Plagiogyriaceae: Plagiogyria semicordata Stygnidae: Nomoclastes quasimodo, Stygnus (MB). MAGNOLlOPHYTA. Magno1iop-sida. gertschi (PI). Hexapoda. Coleoptera: Asterales. Asteraceae: Montanoa ovalifolia Erirhinidae: Lissorhoptrus persimilis (OB); subsp. ovalifolia (FU). Euphor-biales. Staphylinidae: Pseudopsis grossa (I-IE). I-Ialoragales. Gunneraceae: Gunnera saint- I-Iemiptera. Miridae: Rhinacloa incaicus (SV). johnii (MD), Myrtales. Onagraceae: Fuchsia I-Iymenoptera. Pompilidae: Pompilocalus petiolaris, F. venusta (BR). Sapindales. ruminahui, P. vinicolor (RJ). Lepidoptera. Zygophyllaceae: Bulnesia carrapo (HI-I), Nymphalidae: Actinote des miala (AD). Urticales. Cecropiaceae: Cecropia goudotiana Trichoptera. Polycentropidae: Polycentropus (FR). Violales. Passifloraceae: Passiflora cuspidatus (I-IL). VERTEBRAT A. adulterina, P. cuatrecasasii, P. lanata (ES). Actinopterygii. Characifonnes. Characidae: ARTHROPODA. Arachnida. Opiliones. Creagrutus brevipinnis, C. caucanus (I-IV). Stygnidae: Eutimesius ornatus, Metaphareus Amphibia. Anura. I-Iylidae: Gastrotheca albimanum, Phareus raptator (PI). I-Iexapoda. argenteovirens (DL). Aves. Apodiformes. Curculionidae: Ecnomo-rhinus quasimodus Apodidae: Cyp selo ides lemosi (MU). (VN). I-Iemiptera. Enicocephalidae: Craciformes. Cracidae: Ortalis g. guttata Enicocephalus sturmi (GT). Lepidoptera. (MU). Piciformes. Picidae: Picumnus Nymphalidae: Actinote anteas, A. callianthe, granadensis (M U). Tinamiformes. Tinamidae: Actinotee. equatoria,A. eresia,A. hylonome, Nothocercus julius (MU). Mamma1ia. A. inaequalis, A. mel amp epl.os (AD). Artiodactyla. Cervidae: Pudu mephistophi/es VERTEBRAT A. Aves. Columbiformes. (MU). Carnivora. Procyonidae: Nasuella Columbidae: Columbina passerina parvula

60 Juan J. Morrone

(HG). Coraciformes. Momotidae: Momotus montane forests ecoregion momota (HG). Passeriformes. Certhiidae: (Dinerstein et al., 1995: 97). Campylorhynchus griseus (MU). Marnrnalia. Taxa. MAGNOLIOPHYT A. Magnoliop-sida. Lagomorpha. Leporidae: Sylvilagus Ari sto loch iales. Aristo loch iaceae: jloridanus purgatus (M U). Primates. Aristolochia goudotii, A. nummularifolia, A. Callithrichidae: Saguinus leucopus (EM). p annosoides (GO). Fabales. Fabaceae: Rodentia. Echimyidae: Proechimys Centrosema tetragonolobum (SW). magdalenae (PT); Sciuridae: Sciurus ARTHROPODA. Crustacea. Decapoda. granatensis norosiensis (HG). Trichodactylidae: Forsteria venezuelensis (RO, ML). Hexapoda. Coleoptera. Venezuela n Llanos province (Fig. 3.7) Curculionidae: Loncophorus santaros ae, Great part ofVenezuela and northeastern Co- Naupactus jolyi, N. sanfilippoi, Tyloderma lombia. brevisquameum, T. variabile (80, CK, WR). Synonyrns. Sabana district (Cabrera & Yepes, Lepidoptera. Nymphalidae: Actinote amida 1940: 14), Caquetio province (Fittkau, 1969: (AD). VERTEBRATA. Actinopterygii. 642), Sabana province (Cabrera & Willink, Characiformes. Curimatidae: Cyphocharax 1973: 63), Caribbean centre (Müller, 1973: 57), meniscaprorus, Steindachnerina pupula (VS, Orinoqu ia province (Ringuelet, 1975: 107), Lla- VT). Crocodylia. Crocodylidae: Crocodylus nos province (Rivas-Martínez & Navarro, intermedius (DO). 1994: map; Morrone, 1999: 5), Septentrional Venezuelan province (Rivas-Martínez & Na- Western Ecuador province (Fig. 3.8) varro, 1994: m ap), Llanos ecoregion Western Ecuador and southwestern Colom- (Dinerstein et al., 1995: 104), Llanos dry forests bia. ecoregion (Dinerstein et al., 1995: 100), and Synonyrns. Guayas province (Ringuelet, 1975:

3::..",' ;0 c)" ••

Fig. 3. South American biogeographic provinces ofthe Caribbean subregion. 1, Maracaibo; 2, Coastal Venezuela; 3, Trinidad and Tobago; 4, Chocó; 5, Cauca; 6, Magdalena; 7, Venezuelan Llanos; 8, Western Ecuador; 9, Dry Ecuador; l O, Galapagos islands; 11, Tumbes-Piura.

61 Toward a c1adistic model for the Caribbcan subregion 107), Western Ecuador moist forests ecoregion C. sandrae, Neozimiris pinta, N. pinzon, (Dinerstein et al., 1995: 96), and Western Poecilochr oa bifasciata, Trachyzelotes Ecuadorian province (Morrone, 1999: 6). kulckrynskii, Zelotes reformans (BA). Taxa. MAGNOLlOPHYT A. Magnoliop-sida. Chilopoda. Geophilomorpha. Chileno-philidae: Myrtales. Onagraceae: Fuchsia macrostigma, Pachymerium pereirai (SP); Geophilidae: F. polyantha, F. sylvatica (BR). Pachymerium pereirai (PE); Schendylidae: VERTEBRA TA. Actinopterygii. Characi-for- Nannopodellus purpurascens, Nesondyla mes. Curimatidae: Pseudocurimata boehlcki, nealota, Pectiniunguis albermarlensis, P. P. boulengeri, P. patiae (VI). Amphibia. krausi (PE). Scolopen-dromorpha. Anura. Hylidae: Gastrotheca angustifrons Cryptotidae: Cryptops beeb ei (SP); (DL). Scolopendridae: Scolopendra gal ap a- goensis (SP). Crustacea. Anostraca. Dry Ecuador province (Fig. 3.9) Thamnocephalidae: Dendrocephalus Western Ecuador. sarmentosus (CQ). Ostracoda. Cypridinidae: Synonyms. Guayas province (Ringuelet, 1975: Skotsbergia galapagensis (KI); Thaumato- 107), Ecuadorian province (Rivas-Martínez & cyprididae: Danielopolina styx (KI). Navarro, 1994: map), Ecuadorian dry forests Hexapoda. Coleoptera. Anobiidae: Blairinus ecoregion (Dinerstein et al., 1995: 100), Gua- (PH); Anthribidae: Ormiscus variegatus (PK); yaquil flooded grasslands ecoregion Buprestidae: Chrysobothris williamsi (PK); (Dinerstein et al., 1995: 107), and Dry Ecuador Carabidae: Bembidion galap agoensis, province (Morrone, 1999: 6). Calosoma galapageium, C. granatense, C. Taxa. ARTHROPODA. Hexapoda. Coleop- leleuporum, C. linelli, Cicindela galapa- tera. Curculionidae Galapaganus femoratus goensis, Selenophorus obscuricornis (DS, species group, G. propinquus (LN). Diptera. DT, DU, PH); Cerambycidae: Nesoeme(PH); Ditomyiidae: Rhipidita pleciodes (AM). Chrysomelidae: Docema and Scutobruchus Hymenoptera. Apidae: Geotrigona ceratioborus (PK); Cleridae: Pelonium leucogastra (CD). VERTEBRATA. Ma- longfildae (PK); Coccinelidae: Diomus rnrnalia. Rodentia. Echimyidae: Proechimys galapagoensis (PK); Cucujidae: Catharto- decumanus (PT). silvanus tropicalis (PK); Curculionidae: Anchonus galapagoensis, Galapaganus GaIapagos islands province (Fig. 3.10) spp., Lembodes subcostatus, Colón archipelago, in the Pacific Ocean, 950 Neopentarthrum mutchleri, Pagiocerus km west from the coast of Ecuador, which frontalis{LN, PK); Dytiscidae: Copelatus comprises 15 larger islands, as well as several galap agoensis, Thermoncetes basillaris smaller islands. galapagoensis (PK); Elateridae: Conoderus Synonyrns. Galapagos islands province (Ca- galap agoensis (PK); Hydrophilidae: brera & Willink, 1973: 46; Rivas-Martínez & Galapagodacnum, Enochrus ob scurus, Navarro, 1994: map; Morrone, 1999: 5), Tropisternus lateralis (PK); Mordellidae: Galapagos islands centre (Müller, 1973: 106), Mordellistena galap agoensis (PK); and Galapagos islands xeric scrub ecoregion Mycetophagidae: Litargus balteatus (PK); (Dinerstein et al., 1995: 111). Nitidulidae: Acribus (PH, PK); Phalacridae: Taxa. MAGNOLlOPHYTA. Magnoliop-sida. Phalacrus darwini (PK); Scarabeidae: Caryophyllales. Cactaceae: Opuntia Nesoryctes (PH); Rhizophagidae: Bactridium galapageia (CW). ARTHROPODA. Arach- insularis (PK); Staphylinidae: Neolinus, nida. Araneae. Gnaphosidae: Camillina Pinostygus (CU, PH); Tenebrio-nidae: galapagoensis, C. isabela, C. pecki, C. isla, Blapstinus, Pedoneces, Stomion (AB, PH);

62 Juan J. Morrone Trogidae: Omorgus indigenus (SC). Diptera. Stygnidae: Stygnus mediocris (PI). Dolichopodidae: Amblypsilopus depilis, VERTEBRA TA.Actinop-terygii Characiformes. Asyndetus bursericola, A. cavagnaroi, A. Curimatidae: Pseudo-curimata peruana (VR). maelfaiti, A. versicolor, Chrysotus baerti, C. Squamata. Vipe-ridae: Bothrops barnetti (CV). brevicornis, Cymatopus setosus, Medereta Aves. Passeri-formes. Corvidae: Cyanocorax galapagensis, Thinophilus hardyi (BS). mystacalis (MU); Fringillidae: Carduelis Hemiptera. Miridae: Rhinacloa insularis, R. siemiradzkii, Piezorhina cinerea, Saltator longirostris, R. mella, R. rubescens, R. albicollis, Sicalis taczanowskii (SI). Mammalia. usingeri (SV). Hymenoptera. Formicidae: Rodentia. Echimyidae: Proechimys decumanus Strumigenys longispinosa, S. marginiventris species group, P. rosa (PT). (LG). Neuroptera. Hemerobiidae: Megalomus darwini (MN). Orthoptera. Acrididae: ACKNOWLEDGMENTS Halmenus spp. (CI). VERTEBRATA. Chelonia. Testudinidae: Chelonoidis 1 arn grateful to Dalton de Souza Arnorim, elaphantopus (MU). Squamata. Gekkonidae: Mercedes Azpelicueta, David Espinosa, Osear Phyllodactylus spp. (MU); Iguanidae: Flores, Pilar Franco, Irene Goyenechea, Analía Amblyrhynchus cr istatus, Tropidurus Lanteri, Livia León, Jorge Llorente, Rafael albermarlensis (MU, ST). Aves. Ciconiifor- Miranda, Luis Pereira, and Jorge Williams for mes. Laridae: Creagrus furcatus, Larus assistance in discussing many items and/or ful iginosus (MU); Phalacrocoracidae: providing relevant !iterature; and to Jorge Phalacrocorax harrisi (MU); Spheniscidae: Llorente and an anonymous reviewer for Spheniscus mendiculus (MU). Passeriformes. helpful criticisms. Economic support by the Fringillidae: Camarhynchus crassirostris, C. National Geographic grant 6590-99 and heliobathes, C. pallidus, C. parvulus, C. DGAPA-Papiit IN205799 (UNAM) isgratefuIly pauper, C. psittacula, Certhidae olivacea, acknowledged. Geospiza conirostris, G. difficilis, G. fortis, G. fuliginosa, G. magnirostris, G. scandens LITERATURE CITEO (MU); Sturnidae: Nesomimus macdonaldi, N. melanotis, N. parvulus, N. trifasciatus (MU). AALBU,R. L. & C. A. TRIPLEHORN.1991. Mammalia. Carnivora. Otariidae: Arctoce- Pedonoeces G. R. Waterhouse = Blapstinus phalus galapagoensis (MU). Rodentia. Sturm, relevant name changes for California Muridae: Oryzomys galapagoensis (NW). and Galapagos islands species and new insular species from Mexico (Coleoptera: Turnbes-Piura province(Fig. 3.11) Tenebrinidae). Cols. Bull. 45: 169-175. Southern Ecuador and northern Peru. ALBERICO,M. & E. VELASCO.1994. Extended Synonyms. Andean Pacific centre (Müller, description of Platyrrhinus chocoensis 1973: 100),Ecuadorian subcentre (Müller, 1973: from the Pacific lowlands of Colombia. 101), Guayas province (Ringuelet, 1975: 107), Trianea 5: 343-351. Tumbes/ Piura dry forests ecoregion ÁLVAREZ-CASTAÑEDA,S. T., C. A. SALlNAS- (Dinerstein et al., 1995: 100), and Tumbes- ZAVALA& F. DELACHICA.1995. Análisis Piura province (Morrone, 1999: 6). biogeográfico del noroeste de México con Taxa. MAGNOLlOPHYT A. Magnoliop-sida. énfasis en la variación climática y Asterales. Asteraceae: Montanoa ovalifolia mastozoológica. Acta Zool. Mex., n. s. 66: subsp. australis (FU). Myrtales. Onagraceae: 59-86. Fuchsia andrei, F. ayavacensis, F. scherffiana AMORIM,D. S. & M. R. S. PIRES. 1996. (BR). ARTHROPODA. Arachnida. Opiliones. Neotropical biogeography and a method

63 Toward a c1adistic modcl for the Caribbcan subregion for maximum biodiversity estimation. In: Reptilia./n: Goldblatt, P. (ed.), Biological Bicudo, C. E. M. & N. A. Menezes (eds.), relationships between Africa and South Biodiversity in Brazil: A first approach, America Yale University Press, New Haven CNPq, Sao Paulo, pp. 183-219. & London, pp. 244-288. ANDRADE, M. G. 1995. Nymphalidae: BERNAL,H. Y. 1986. Crota/aria (Fabaceae- Acraeinae: Actinote. In: Ruiz, P. M. & P. Faboideae)./n: Pinto, P. & P. M. Ruiz(eds.), Pinto (eds.), Monografías de fauna de Co- Flora de Colombia, monogr. no. 4, Univer- lombia, Instituto de Ciencias Naturales- sidad Nacional de Colombia, Santafé de Bo- Museo de Historia Natural, Santafé de Bo- gotá,pp.I-118. gotá, pp. 1-120. BERRY, P. E. 1982. The systematics and ANDREWS. F. G. 1994. New species of evolution of Fuchsia sect. Fuchsia Metophtha/mus Wollaston from the West (Onagraceae). Ann. Missouri Bot. Gard. 69: Indies (Coleoptera: Lathridiidae: 1-198. Lathridiini). Co/s. Bull. 48: 191-197. BICKEL, D. J. & B. J. SINCLAIR. 1997. The ANDREWS, F. G. 1998. A new species of Dolichopodidae (Diptera) ofthe Galapagos Metophtha/mus Wollaston (Coleoptera: islands, with notes on the New World fau- Latridiidae) from Cuba. Co/s. Bull. 52: 60-62. na. Ent. Scand. 28: 241-270. ARANDA,M., 1. E. ESCOBEDO& C. Pozo. 1997. BORDÓN, C. 1991. El género Macrostylus Registros recientes de Otonyctomys hatti Boheman (Col. Curc. Bracyderinae, (Rodentia: Muridae) en Quintana Roo, Naupactini) en Venezuela. Acta Bio!. México. Acta Zool. Mex., n. s.72: 63-65. Venezuel. 13: 1-50. ARRIAGA, L., C. AGUILAR,D. ESPINOSA& R. BORDÓN,C. 1997. El género Naupactus Dejean JIMÉNEZ (eds). 1997. Regionalización (Coleoptera: Curculionidae) en Venezuela. ecológica y biogeográfica de México. ActaBio!. Venezuel. 17: 11-51. Workshop at the Comisión Nacional para BURKE,H. R. & P. W. KOVARIK.1986. Revision el Conocimiento y Uso de la Biodiversidad of the Neotropical genus Achia Champion (Conabio), November 1997,99 pp. (Coleoptera: Curculionidae). Stud. ASHE,J. S. & S. W. LINGAFELTER.1995. Revision Neotrop. Fauna Environ. 21: 129-168. of Gansia Sharp of Mexico and Central CABRERA,A. & 1. YEPES.1940. Mamíferos sud- America (Coleoptera: Staphy-Iinidae: americanos (vida, costumbres y descrip- Aleocharinae).J. New York Entomol. Soco ción). Historia Natural Ediar, Buenos Ai- 103: 251-280. res. BAERT, L. 1994. The Gnaphosidae of the CABRERA,A. L. & A. WILLlNK. 1973. Biogeo- Galapagos archipelago, their distribution grafla de América Latina. Monografía 13, and the description of the Galapagoan Serie de Biología, OEA, Washington D.C. representatives of the genus Camillina CAMARGO, 1. M. F. & 1. S. MOURE. 1996. Berland. Bull. Inst. Roy. Sci. Nat. Be/gique Meliponini neotropicais: O genero 64: 9-24. Geotrigona Moure, 1943 (Apinae, Apidae, BÁEZ, A. M. 1977. Consideraciones sobre la Hymenoptera), com especial referencia a osteología y el status taxonómico de los filogenia e biogeografia. Arq. Zool., Sao pípidos sudamericanos vivientes Pau/o33: 95-161. (Amphibia, Anura). Physis (Buenos Aires) CAMPBELL, J. A. 1988. The distribution, C,35:321-336. varratrons, natural history, and BAUER,A. M. 1993. African-South American relationships of Porthidium barbouri relationships: A perspective from the (Viperidae). Acta Zoo/. Mex., n. S. 26: 1-32.

64 Juan .r. Morrone CAMPBELL,J. A. 1999. Distribution patterns CLARK,W. E. 1988. Revisionoftheweevilgenus of amphibians in Middle America. In: Loneophorus Chevrolat (Coleoptera: Duellman, W. E. (ed.), Patterns of Curculionidae, Anthonominae). Quaest. distribution of amphibians: A global Entomo/. 24: 465-520. prespective, The John Hopkins CLARK,W. E. 1992.TheAnthonomus marmoratus University Press, Baltimore and London, species group (Coleoptera: Curculionidae). pp, 111-210. Trans.Amer.Entomo!.Soco118:129-145. CAMPBELL,1.M. 1991. Peplomicrus iviei, new CLARK, W. E. 1993. The weevil genus species: First report of the family Neomastix Dietz (Coleoptera: Curculioni- Micropeplidae (Coleoptera) from the West dae, Anthonomini). Cols. Bull. 47: 1-19. Indies. Cols. Bull. 45: 37-41. COIlEN,R. G. 1998. Anostraca./n: Morrone, 1. CAMP13ELL,1. M. & S. B. PECK. 1989. Pinos- 1. & S. Coscarón (eds.), Biodiversidad de tygus galapagoensis, a new genus and artrópodos argentinos: Un enfoque species of eyeless rove beetle (Coeloptera: biotaxonómico, Ediciones Sur, La Plata, pp. Staphylinidae: Paederinae) from a cave in 491-501. the Galapagos islands, Ecuador. Cols. Bull. COSCARÓN, S. 1987. El género Simulium 43: 397-405. Latreille en la región Neotropical: Análisis CAMPOS,M. R. 1997. Primeros registros de la de los grupos supraespecíficos, especies tribu Kingsleyini (Crustacea: Decapoda: que los integran y distribución geográfica Pseudotelphusidae) para Colombia. (Simuliidae, Diptera). Museu Paraense Caldasia 19:269-277. Emilio Goeldi, Belem, pp. 1-111. CARLTON,C. E. 1989. Revision of the genus COSCARÓN,S. & C. L. COSCARÓN-ARIAS.1995. Eutriehites LeConte (Coleoptera: Psela- Distribution of Neotropical Simuliidae phidae). Cols. Bul/. 43: 105-119. . (Insecta, Diptera) and its areas of CARRANZA-GONZÁLEZ,E. 1993. Una nueva va- endemism. Rev. Aead Colombo Ciene. 19: riedad de Styrax argenteus (Styracaceae) 717-732. del NE de Querétaro (México). Acta Bol. COSCARÓN, S., S. IBAÑEZ-BERNAL & c. L. Mex. 23: 35-40. COSCARÓN-ARIAS. 1996. Revisión de CASTILLO,C. & P. REYES-CASTILLO.1984. Bio- Simulium (Psilopelmia) Enderlein en la re- sistemática del género Petrejoides Kuwert gión Neotropical y análisis c1adístico de (Coleoptera, Lamellicornia, Passalidae). sus especies (Diptera: Simuliidae). Acta Acta Zoo/. Mex., n. s. 4: 1-84. Zoo/. Mex., n. S. 69: 37-104. CASTILLO,c., L. E. RIVERA-CERVANTES& P. RE- CRACRAFT, J. 1988. Deep-history biogeo- YES-CASTILLO. 1988. Estudio sobre los graphy: Retrieving the historical pattern of Passalidae (Coleoptera: Lamellicornia) de evolving continental biotas. Syst. Zoo/. 37: la Sierra de Manantlán, Jalisco. Acta Zool. 221-236. Mex., n. s. 30: 1-20. CRACRAFT,1.& R. O. PRUM. 1988. Patterns and CIGLlANO,M. M. & C. E. LANGE.1998. Orthop- processes of diversification: Speciation tera./n: Morrone, 1. 1. & S. Coscarón (eds.), and historical congruence in sorne Biodiversidad de artrópodos argentinos: Neotropical birds. Evolution 42: 603-620. Un enfoque biotaxonómico, Ediciones Sur, CRAW,R. C., 1. R. GREHAN& M. 1. HEADS.1999. La Plata, pp. 67-83. Panbiogeography: Traeking the history CLARK, W. E. 1987. Revision ofthe unipus- of lije. Oxford Biogeography Series 11, tulatus group of the weevil genus Oxford University Press, New York. Anthon omus Germar (Coleoptera: CRAW, R. C. & R. D. M. PAGE. 1988. Curculionidae). Cols. Bull. 41: 73-88. Panbiogeography: Method and rnetaphor

65 Toward a cladistic model for the Caribbean subregion in the new biogeography./n: Ho, M. -W. y Bias (eds.), Advances in coleopte-rology, S. W. Fox (eds), Evolutionary processes European Association ofColeop-terology, and metaphors, John Wiley and Sons, pp. Torino, pp. 193-200. 163-189. DESENDER,K., L. BAERT& J.-P. MAELFAIT.1992a. CROlZAT,L. 1952. Manualofphytogeography. Distribution and speciation of carabid beetles Junks Gravenhage, The Hague. in the Galápagos archipelago (Ecuador). Bull. CROIZAT,L. 1976. Biogeografla analítica y sin- lnst. Roy. Sci. Nat. Belgique62: 57-65. tética ('panbiogeografia') de las Améri- DESENDER,K., L. BAERT& J.-P. MAELFAIT.1992b. cas. Biblioteca de la Academia de Ciencias El N iño-events and the establishment of Físicas, Matemáticas y Naturales, Caracas. ground beetles in the Galápagos CROTHER,B. 1. & c. GUYER. 1996. Caribbean archipelago. Bull. /nst. Roy. Sci. Nat. historical biogeography: Was the dispersal- Belgique 62: 67-74. vicariance debate eliminated by an DINERsTEIN,E. D. M. OLSON,D. J. GRAIIAM,A. extraterrestrial bolide? Herpetologica 52: L. WEBSTER,S. A. PRIMM,M. P. BOOKBINDER 440-465. & G. LEDEC.1995. Una evaluación del es- CROTIlER, B. l., M. M. MIYAMOTO& W. F. tado de conservación de las eco-regiones FRESCII. 1986. Phylogeny and terrestres de América Latina y el Caribe. biogeography of the lizard family World Bank, Washington, D.C., 135 pp. Xantusiidae. Syst. Zool. 35: 37-45. DONoso-BARRos, R. 1965. Contribución al co- DELGADO,L. & J. BLACKALLER-BAGES.1997. A nocimiento de los cocodrilos de Venezue- new Mexican species of Homoiosternus la. Physis (Buenos Aires) 25: 387-400. (Coleoptera: Melolonthidae; Rutelinae) . .J. DONoso-BARROS, R. 1966. Contribución al co- New York Entomol. Soco 105: 170-179. nocimiento de los cocodrilos de Venezue- DELGADO,L. & R. HERNÁNDEZ.1998. Anopsiostes la. Continuación. Physis (Buenos Aires) 26: pauliani new species and a first record of 15-32. the genus from Mexico (Coleop-tera: DORADO,O. & D. M. ARIAS.1992. Brongniartia Scarabaeidae: Ceratocanthinae). J New York montalvoana (Fabaceae: Faboideae), una Entomo!. Soco106: 105-108. especie nueva de la cuenca del Balsas. Acta DELOYA,C. 1995. A new species of Cotinis Bol. Mex. 17: 13-17. (Cotinis) Burmeister from Mexico DUELLMAN,W. E. 1989. Lista anotada y clave (Coleoptera: Melolonthidae, Cetoninae). de los sapos marsupiales (Anura: Hylidae: Cols. Bu!!' 49: 149-152. Gastrotheca) de Colombia. Caldasia 16: DELOYA,C. & B. C. RATCLIFFE.1988. Las espe- 105-111. cies de Cotinis Burmeister en México EMMoNs, L. H. 1990. Neotropical rairforest (Coleoptera: Melolonthidae: Cetoniinae). mammals: Ajield guide. The University of Acta Zool. Mex., n. s. 28: 1-52. Chicago Press, Chicago & London. DEMANGE, J .-M. & L. A. PEREIRA. 1985. ESCALANTE,P., A. G. NAVARRO& A. T. PETERSON. Géophilomorphes (Myriapoda, Chilopoda) 1998. Un análisis geográfico, ecológico e de la Guade loupe et ses dépendances. histórico de la diversidad de aves terres- Bul!. Mus. Natn. Hist. Nat., Paris, 4e. sér. tres de México. In: Rarnarnoorthy, T. P., R. 7: 181-199. Bye, A. Lot& J. Fa(eds.), Diversidad bio- DESENDER,K., L. BAERT& J.-P. MAELFAIT.1991. lógica de México: Orígenes y distribución, Evolutionary systematics of Calosoma Instituto de Biología, UNAM, Mexico, Weber carabid beetles of the Galápagos D.F., pp. 279-304. archipelago, Ecuador (Coleop-tera: ESCOBAR,L. K. 1988. Passitloraceae: Passi-flo- Carabidae)./n: Zunino, M., X. Bel1és & M. ra, subgéneros T'acsoni a, Rathea,

66 Juan J. Morrone

Manicata y Distephana.ln: Pinto, P. & G. GENIER,F. & H. F. HOWDEN.1999. Two new Lozano (eds.), Flora de Colombia, monogr. Central American Onthophagus Latreille of no. 10, Universidad Nacional de Colombia, the mirabilis species group (Coleoptera: Santafé de Bogotá, pp. 1-138. Scarabaeidae, Scarabaeinae). Cols. Bul!. 53: FERNÁNDEZ,A. M., D. RODRÍGUFZ& V. HERNÁNDEZ- 130-144. ORTlZ. 1997. Notas sobre el género GIESBERT, E. F. 1993. New species of Anastrephas Schiner en Cuba con descrip- Ceram bycidae (Coleoptera: Cerambycidae) ción de una nueva especie (Diptera: from Guatemala and Chiapas, Mexico. Cols. Tephritidae). Folia Entomol. Mex. 99: 29-36. Bul!. 47: 137-149. FERNÁNDEZ,1.L. & A. COGOLLO.1992. Notas GONZÁLEZ,F. A. 1990. Aristolochiaceae. In: sobre Matisia longiflor a Gleas. Rangel, J. O., A. Cadena, G. Correal & R. (Bombacaceae). Caldasia 17: 173-176. Bernal (eds.), Flora de Colombia, monogr. FERNÁNDEZ-ALONSO,J. L. 1997. Nueva especie no. 12, Universidad Nacional de Colombia, de Condalia Cavo (Rhamnaceae) y notas Santafé de Bogotá, pp. 1-184. sobre los géneros de la familia en la flora GONZÁLEZ-EuzONDO,M. S. & M. GONZÁLEZ- de Colombia. Caldasia 19: 10 1-108. EUZONDO. 1992. Una nueva especie de FUNT, O. S. Jr. 1996. Studies ofNeotropical Arbutus (Ericaceae, Arbuteae) de la Sierra caddisflies LV: Trichoptera ofTrinidad and Madre Occidental, México. Acta Bol. Mex. Tobago. Trans. Amer. Entomol. Soco 122: 17:7-12. 67-113. GORDON, R. D. 1977. Classification and FITTKAU, E. J. 1969. The fauna of South phylogeny of the New World America. In: Fittkau, E. J. J. IIIies, H. Sticholodinae (Coccinellidae). Coleopts. Klinge, G. H. Schwabe & H. Sioli (eds.), Bul!. 31: 185-228. Biogeography and ecology in South GORDON,R. D. 1982. West Indian Coccinellidae América, 2, Junk, The Hague, pp. 624-650. 111 (Coleoptera): A new species of FORERO,E., E. CARBONO,C. I. OROZCO,E. ORTEGA, Nexophallus Gordon from Trinidad. J. E. RAMOS,R. RUIZ,O. SALAZARDEBENAVIDES Coleopts. Bul!. 36: 118-120. & L. A. VIDAL.1983. Connaraceae.ln: Pinto, GORDON, R. D. 1994a. West Indian P. & P. M. Ruiz (eds.), Flora de Colombia, Coccinellidae V (Coleoptera): A review of monogr. no. 2, Universidad Nacional de Co- Coccidulini and additions to Psorolyma lombia, Santafé de Bogotá, pp. 1-83. Sicard.J New York Entomol. Soco 102: 222- FRANCO,P. & c. c. BERG.1997. Distributional 231. patterns of Cecropia (Cecropiaceae): A GORDON, R. D. 1994b. West Indian panbiogeographic analysis. Caldasia 19: Coccinellidae VI (Coleoptera): New genera 285-296. and species od Sticholotidini and a FRANK,1.H. 1981. A revision ofthe New World cladistic analysis of incIuded genera. J species of the genus Neob isnus New York Entomol. Soco 102: 232-241. Ganglbauer (Coleoptera: Staphylin idae: GRIMALDI,D. A. 1988. RelictsinDrosophilidae Staphylininae). Occas. Papo Florida State (Diptera).ln: Liebherr,J. K. (ed.),Zoogeography Col/o Arthrop. 1: 1-58. of Caribbean , Cornell University FREIRE,S. E. 1993. A revision of Chionolaena Press, Ithaca& London, pp, 183-213. (Compositae, Gnaphalieae). Ann. Missouri GRIMALDI,D., C. MICHALSKI& K. SCIlMlDT. Bot. Gard 80: 397-438. 1993. Amber fossil Enicocephalidae FUNK, V. A. 1982. The systematics of Mon- (Heteroptera) from the Lower Cretaceous tanoa (Asteraceae, Heliantheae). Mem. of lebanon and Oligo-Miocene of the New York Bot. Gard 36: 1-133. Dominican Republic, with biogeographic

67 Toward a c\adistic model for (he Caribbean subrcgion

analysis of Enicocephalus. Am. Mus. HEDGES,S. B., C. A. HASS& L. R. Mxxson. Novit. 3071: 1-30. 1992. Caribbean biogeography: Molecular GUTIÉRREZ,E. 1995. Annotated checklist of evidence for dispersal in West Indian Cuban cockroaches. Trans. Amer. Entomol. terrestrial vertebrates. Proc. Natl. Acad. Soc.121:65-84. Sci. USA 89: 1909-1913. GUYER, C. & 1. M. SAVAGE. 1986. Cladistic HERMAN,L. H. 1972. Revision of Bledius and relationships among anoles (Sauria: related genera. Part 1. The aequatorialis, Iguanidae). Syst. Zoo/. 35: 509-531. mandibularis, and semiferrugineus groups HALFfTER,G. 1978. Un nuevo patrón de dis- and two new genera (Coleoptera, persión en la zona de transición mexicana: Staphylinidae, Oxytelinae). Bull. Am. Mus. El mesoarnericano de montaña. Folia Nat. Hist. 149: 113-254. Entomol. Mex. 39-40: 219-222. HFRMAN,L. H. 1975. Revision and phylogeny HALFFTER, G. 1987. Biogeography of the of the monogeneric subfam ily montane entornofauna ofMexico and Cen- Pseudopsinae for the world (Staphylinidae, tral Arnerica. Annu. Rev. Entomol. 32: 95- Coleoptera). Bu!!'Amer. Mus. Nat. Hist. 155: 114. 243-317. HAMILTON,R. W. 1994. Revision ofthe New HERMAN,L. H. 1986. Revision of Bledius. Part World genus Pilolabus Jekel (Coleoptera: IV. Classification of species groups, Attelabidae). Trans. Amer. Entomol. Soco phylogeny, natural history, and catalogue 120: 369-411. (Coleoptera, Staphylinidae, Oxytelinae). HAMILTON, R. W. 1997. A new species of Bull. Am. Mus. Nat. Hist. 184: 1-367. Euscelus Schoenherr frorn Mexico HERMAN,L. H. 1991. Revision ofthe subtribe (Coleoptera: Attelabidae). Cols. Bull. 51: Cylindroxystina (Coleoptera: Staphyli- 364-370. nidae: Paederinae). Bull. Amer. Mus. Nat. HAMILTON, S. W. 1988. Historical Hist. 203: 1-83. biogeography oftwo groups ofCaribbean HERNÁNDEZ,1., A. HURTADO,R. ORTIZ & T. Polycentropus (Trichoptera: WALSCH13URGER.1992a. Unidades biogeo- Polycentropidae).ln: Liebherr, 1. K. (ed.), gráficas de Colombia. In: Halffter, G. (ed.), Zoogeography of Caribbean insects, La diversidad biológica de lberoamérica, Cornell University Press, Ithaca & London, Acta Zoológica Mexicana, Vol. Esp. 1992, pp. 153-182. Cyted-D, Programa Iberoamericano de HAROLD,A. S. & R. P. V ARI.1994. Systematics Ciencia y Tecnología para el Desarrollo, of the Trans-Andean species of Instituto de Ecología, A.C., Xalapa, pp. Creagrutus (Ostariophysi: Characidorrnes: 105-l5l. Characi-dae). Smithson. Contrib. Zoo/. 551 : HERNÁNDEZ,1., A. HURTADO,R. ORTIZ & T. 1-31. WALSCH13URGER. 1992b. Centros de HEDGES,S. B. 1996a. Historical biogeography endemismo en Colombia. In: Halffter, G. of West Indian vertebrates. Annu. Rev. (ed.), La diversidad biológica de Eco!. Sysl. 27: 163-196. Iberoarnérica, Acta Zoológica Mexicana, HEDGES,S. B. 1996b. The origin ofWest Indian Vol. Esp. 1992, Cyted-D, Programa Ibero- arnphibians and reptiles. In: Powell, R. & americano de Ciencia y Tecnología para el R. W. Henderson (eds.), Contributions to Desarrollo, Instituto de Ecología, A.C., west Indian herpetology: A tribute to Albert Xalapa, pp. 175-190. Schwartz, Society for the Study of HERNÁNDEZ,1., T. W ALSCH13URGER,R. ORTIZ& Amphibians and Reptiles, Ithaca, pp. 95- A. HURTADO.1992c. Origen y distribución 128. de la biota suramericana y colombiana. In:

68 Juan J. Morrone Halffter, G. (ed.), La diversidad biológica métodos. In: Métodos cuantitativos en la de Iberoamérica, Acta Zoológica Mexica- biogeografía, Publ. 12, Instituto de na, Vol. Esp. 1992, Cyted-D, Programa Ibe- Ecología, A. c., Xalapa, pp. 41-120. roamericano de Ciencia y Tecnología para KORNICKER, L. S. & T. M. IUFFE. 1989. el Desarrollo, Instituto de Ecología, A.C., Troglobitic Ostracoda (Myodocopa: Xalapa, pp. 55-104. Cypridinidae, Thaurnatocyprididae) frorn HESPENIIEIDE,H. A. 1997. New species and a anchialine pools on Santa Cruz island, new nam e fro Antillean Buprestidae Galapagos islands. Smithson. Contrib. (Coleoptera). J. New York. Entamo!. Soco Zool. 483: 1-38. 105: 193-198. LANTERI,A. A. 1992. Systernatics, cladistics HOWDEN,A. T. 1992. Review ofthe New World and biogeography of a new weevil genus, eyeless weevils with uncinate tibiae Galapaganus (Coleoptera: Curculionidae) (Coleoptera, Curculionidae: Molytinae, from the Galapagos islands, and coasts of Cryptorhynchinae, Cossoninae). Mem. Ecuador and Peru. Trans. Amer. Entamo!. Entomol. Soco Canada 162: 1-76. Soco 118: 227-267. HOWDEN,H. F. 1988. A new genus and fournew LATTKE, J. & W. GOITÍA. 1997. El género species ofNew World Trichiinae (Coleoptera: Strumigenys (Hyrnenoptera: Formicidae) Scarabaeidae). Ca/s. Bu!!' 42: 241-250. en Venezuela. Caldasia 19: 367-396. HOWDEN, H. F. 1994a. A new Geotrupes LIEBHERR,J. K. 1986. Barylaus, new genus Latreille (Coleoptera: Scarabaeidae) from (Coleoptera: Carabidae) edemic to the West the Sierra de las Minas range of Guatema- Indies with Old World affinities. J. New la. Cols. Bu/!. 48: 345-348. York Entamo!. Soco 94: 83-97. HOWDEN, H. F. 1994b. A new Apeltastes LIEI3I1ERR,1.K. 1988a. The Caribbean: Fertile Howden and notes on Guatemalan ground for zoogeography. In: Liebherr, J. Paragnorimus Becker (Scarabaeidae: K. (ed.), Zoogeography of Caribbean Trichiinae). Cols. Bu/I. 48: 349-353. insects, Cornell University Press, Ithaca & IVIE,M. A.& D. S. SIKES.1995. Description of London, pp. 1-14. Micromyrmex asclepia, new species, with LIEBHERR,1.K. 1988b. Biogeographic patterns a redefinition ofthe genus and life history of West Indian carabid beetles notes (Curculionidae: Otidocephalina). (Coleoptera). In: Liebherr, 1. K. (ed.), Ca/s. Bull. 49: 235-248. Zoogeography of Caribbean insects, JAMESON,M. L. 1990. Revision, phylogeny and Cornell University Press, Ithaca & London, biogeography ofthe genera Parabyrsopolis pp. 121-152. Ohaus and Viridimicus, new genus LIEBIIERR,1. K. 1988c. Redefinition of the (Coleoptera: Scarabaeidae: Rutelinae). Cols. supertribe Odacantithae, and revision of Bu/!. 44: 377-422. the West Indian Lachnophorini KEVAN, D. K. M. 1977. The American (Coleoptera: Carabidae). Quaest. Entomol. Pyrgornorphidae (Orthoptera). Rev. Soco 24: 1-42. Entamo!. Argent. 36: 3-28. LIEBHERR,J. K. 1988d. General patterns in West KLUGE,A. G. 1988. Parsimony in vicariance Indian insects, and graphical biogeographic biogeography: A quantitative rnethod and analysis of sorne circum-Caribbean a Great Antillean example, Syst. Zool. 37: Platynus beetles (Carabidae). Syst. Zool. 37: 315-328. 385409. KOIll.MAN, B. & S. SÁNCHEZ.1984. Estudio LIEBIIERR,J. K. 1997. Review of Antillean areográfico del género Bursera Jacq. Ex L. G/ypto/enus Bates (Coleoptera: Carabidae), (Burseraceae) en México: Una síntesis de with description of a new species

69 Toward a cladistic model for the Caribbean subregion precinctive to St. Vincent. Stud. Neotrop. Contributions to West Indian herpetology: Fauna Environ. 32: 89-99. A tribute to Albert Schwartz, Society for LIZARRALDEDEGROSSO,M. 1982. Redescripción the Study of Amphibians and Reptiles, del género Helaeomyia Cresson y reivin- Ithaca, pp. 141-155. dicación de Mimapsilopa Cresson MAAS, P. 1. M. & H. MAAS-VAN DE KAMER. (Diptera, Ephydridae). Physis (Buenos Ai- 1988. Burmaniaceae.ln: Pinto, P. & G. Lo- res) C, 40: 121-128. zano (eds.), Flora de Co 10mbia, monogr. no. LLORENTE,J. E. 1996. Biogeografía de artrópo- 7, Universidad Nacional de Colombia, dos de México: ¿Hacia un nuevo enfoque? Santafé de Bogotá, pp. 33-124. In: Llorente, J. E., A. N. García & E. MAcF ADDEN,B. 1. 1980. Rafting mammals or González (eds.), Biodiversidad, taxonomía drifting continents?: Biogeography of the y biogeografía de artrópodos de México: Greater Antilles insectivores Nesophontes Hacia una síntesis de su conocimiento, and Solenodon. 1. Biogeogr. 7: 11-22. UNAM, México, D.F., pp. 41-56. MAGALIIAES, C. & M. TÜRKAY. 1996a. LLORENTE,1.E., L. OÑATE,A. LUIS& 1. VARGAS. of the Neotropical freshwater 1997. Pap ilion idae y Pieridae de México: crab family Trichodactylidae 1. The generic Distribución geográfica e ilustración. Co- system with description of sorne new ge- misión Nacional para el Conocimiento y nera (Crustacea: Decapoda: Brachyura). Uso de la Biodiversidad (Conabio) y Fa- Senckerberg. Biol. 75: 3-95. cultad de Ciencias, UNAM, México, D.F. MAGALIIAES,C. & M. TÜRKAY.1996b. Taxonomy LOBO,]. M. & G. HALFFTER.1994. Relaciones en- of the Neotropical freshwater crab family tre escarabajos (Coleoptera: Scara-baeidae) Trichodactylidae 11. The genera Forsteria, y nidos de tuza (Rodentia: Geomyidae): Melocarcinus, Sytviocarcinus, and Implicaciones biológicas y biogeográficas. Zilch.iop sis (Crustacea: Decapoda: ActaZool. Mex., n. s. 62: 1-9. Brachyura). Senckerberg Biol. 75: 97-130. LOUREN<;:O,W. R. 1990. A new species of MATIIIS, W. N. 1980. Studies of Ephydrinac Peucetia from Colombia (Araneae, (Diptera: Ephydridae), III: Revisions of Oxyopidae). Caldasia 16: 193-195. sorne Neotropical genera and species. LouREN<;:o,W. R. & E. FLORES.1989. Los es- Smithson. Contrib. Zool. 303: 1-50. corpiones (Chelicerata) de Colombia. 1. La McHuGII,1. V. 1990. Carinisphindus, a new fauna de la isla Gorgona. Aproximación genus and three new species of biogeográfica. Caldasia 16: 66-70. Neotropical Sphindidae (Coleoptera: LYDEARD,C; M. C. WOOTEN& A. MEYER.1995. Clavicornia). Cols. Bull. 44: 307-322. Mo lecu les, morphology, and area MÉNDEZ-LARIOS,1. & 1. L. VII.LASEÑOR.1995. Re- cladograms: A cladistic and biogeographic visión taxonómica del género Teiranema analysis of Gambusia (Teleostei: (Scrophulariaceae). Acta Bot. Mex. 32: 53-68. Poecilidae). Syst. Biol. 44: 221-236. MILLER, W. V. 1995. Two new species of LYNCH,1.0.1982. Relationships ofthe frogs of Heterocerus Fabricius from Mexico the genus Ceratophrys (Leptodactylidae) and (Coleoptera: Heteroceridae). Cols. Bull. their bearing on hypotheses of Pleistocene 49: 249-252. forests refugia in South America and MONSERRAT,V. 1. 1997. Revisión del género punctuated equilibria.Syst. Zool. 31: 166-179. Megalomus de Latinoamérica (Neuroptera, LYNCH,1. D. 1996. The relationships of the Hemerobiidae). Fragm. Entomo!. (Roma) Hispaniolan frogs ofthe subgenus Pelorius 29: 123-206. (Eleutherodactylus: Leptodactylidae). In: MORA,L. E. 1984. Haloragaceae.ln: Pinto, P. Powell, R. & R. W. Henderson (eds.), & P. M. Ruiz (eds.), Flora de Colombia,

70 Juan .J. Morronc monogr. no. 3, Universidad Nacional de Co- (ed.), Zoogeography ofCaribbean insects, lombia, Santafé de Bogotá, pp. 1-178. Cornell University Press, Ithaca & London, MORÓN,M. A. 1995. Review of the Mexican pp.71-120. species of Golofa Hope (Coleoptera: NOWAK,R. M. 1991. Walker 's mammals ofthe Melolonthidae, Dynastinae). Co/s. Bull. 49: World. Fifth edition. Volumes I and 11. The 343-386. John Hopkins University Press, Baltimore MORRONE,J. J. 1994. On the identification of & London, 1629 pp. areas of endernism. Syst. Biol. 43: 438-441. O' BRIEN,C. W. 1996. Two new l.issorhoptrus MORRONE,1. J. 1999. Presentación preliminar rice pests in northern South Arnerica, with de un nuevo esquema biogeográfico de a review of the species in Colombia and América del Sur. Biogeographica 75: 1-16. Venezuela (Coleoptera: Curculionidae). MORRONE,J. J. & M. DELC. COSCARÓN.1996. Trans. Amer. Entomol. Soco 122: 115-134. Distributional patterns of the American O'BRIEN,C. W.&G.1. WIBMER.1989. Revision Peiratinae (Heteroptera: Reduviidae). Zoo!. ofthe Neotropical genus Neohydronomus Mede!. Leiden 70: 1-15. Hustache (Coleoptera: Curculionidae). MORRONE,1. J. & 1. V. CRISCI.1995. Historical Caleapts. Bull. 43: 291-304. biogeography: Introduction to methods. ORTIz-DiAz,1. 1. 1993. Estudio sistemático del Annu. Rev. Eco!. Syst. 26: 373-40 l. género Gouinia (Grarninae, Chloridoideae, MORRONE,1.1., D. ESPINOSA,C. AGUILAR& J. Eragrostideae). Acta Bat. Mex. 23: 1-33. LLORENTE.1999. Preliminary classification OSELLA,G. 1980. Nuovi curculionidi ciechi della ofthe Mexican biogeographic provinces: regione Neotropicale (Coleoptera). Fragm. A parsirnony analysis of endemicity based Entomol. 15: 365-422. on plant, , and bird taxa. Southwest. PAClIECO, V. & B. D. PATTERSON. 1992. Nat. 44: 508-515. Systematics and biogeographic analyses MORRONE, J. J. & E. C. LOPRETTO. 1996. of four species of Sturnira (Chiroptera: Cladistics of the farnily Trichodactylidae Phyl lostom idae), with emphasis on (Crustacea: Decapoda): A reappraisal. .J. Peruvian forms. Mem. Mus. Hist. Nat., U. Comp. Biol. 1: 65-72. N. M S. M (Lima)2l: 57-81. M ÜLLER,P. 1973. The dispersal centres of PAGE,R. D. M. & c. LYDEARD.1994. Towards a terrestrial vertebrates in the Neotropical cladistic biogeography of the Caribbean. rea/m: A study in the evolution of the Cladistics 10: 21-41. Neotropica/ b iot a and its native PAPENFUSS,T. 1. & D. B. WME. 1987. Two new /andscapes. Junk, The Hague. species of plethodontid salamanders MUImLo, M. T. 1988. Pteridophyta- 1. In: (genus Natatritan) from Mexico. Acta Zoo!. Pinto, P. & G. Lozano (eds.), Flora de Co- Mexicana, n. s. 21: 1-16. lombia, monogr. no. 9, Universidad Na- PARENTI, L. R. 1981. A phylogenetic and cional de Colombia, Santafé de Bogotá, biogeographic analysis of cyprinodontiform pp. 1-54. fishes (Teleostei, Atherinomorpha). Bull. NAVARRETE-HEREDIA,1. L. 1997. Descripción Am. Mus. Nat. Hist. 168: 341-557. de Styngetus adrianae sp. nov., incluyen- PARENTI,L. R. 1991. Ocean basins and the do nuevos datos de Styngetus de México biogeography of freshwater fishes. A ustr. (Coleoptera: Staphylinidae). Folia Syst. Bat. 4: 137-149. Entomo!. Mex. 101: 59-71. PATTON,J. L. 1987. Species groups of spiny NICIlOLS, S. W. 1988. Kaleidoscopic rats, genus Proechimys (Rodentia: biogeography of West Indian Scaritinae Echimyidae). Fieldiana, Zoo!. (N. S.) 39: (Coleoptera: Carabidae).ln: Liebherr, 1. K. 305-34-5.

71 Toward a cladistic model for the Caribbean subrcgion

PECK,S. B. 1991. Beetle (Coleoptera) faunas of Indian terestrial vertebrates. Syst. Zool. 30: tropical oceanic islands: With emphasis on 147-155. the Galápagos archipelago, Ecuador. In: RAMÍREZ-GARCÍA,E. & V. HERNÁNDEZ-ORTIZ. Zunino, M., X. Bellés & M. BIas (eds.), 1994. Revisión de la fam ilia Ropalomeridae Advances in coleopterology, European (Diptera) en México. Acta Zool. Mex., n. s. Association of Coleopterology, Torino, pp. 61: 57-85. 177-192. RAMOS, J. A. 1988. Zoogeography of the PECK,S. B. & 1. KUKALOVÁ-PECK.1995. Origin Auchenorhynchous Homoptera of the and biogeography of the beetles Greater Antilles (Hemiptera).ln: Liebherr, (Coleoptera) ofthe Galapagos archipelago, 1. K. (ed.), Zoogeography of Caribbean Ecuador. Can. J Zool. 68: 1617-1638. insects, Cornell University Press, Ithaca & PEREIRA, L. A. & J.-M. DEMANGE. 1997. London, pp. 61-70. Nouvelle contribution a la connaissance RANGEL,1. O., M. AGUILAR,H. SÁNCIIEZ& P. du genre Ctenophilus Cook, 1896, a Lowv. 1995a. Región Costa Pacífica. In: répartition géographique disjointe Rangel, 1. O. (ed.), Colombia: Diversidad (Myriapoda, Chilopoda, Geophilomorpha, biótica 1, Instituto de Ciencias Naturales, Schendyli-dae). Zoosystema 19: 293-326. Convenio Inderena-Universidad Nacional de PEREIRA,L. A., D. FODDAI& A. MINELL!.1997. Colombia, Santafé de Bogotá, pp. 121-139. Zoogeographical aspects of Neotropical RANGEL,1. O., A. GARZÓN& P. Lowv. 1995b. Geophilomorpha (Chilopoda). Ent. Scand. Sierra Nevada de Santa Marta-Colombia 51: 77-86. (con énfasis en la parte norte Transecto PÉREZ, D. E., G. O. DOMINICI, B. HIERRO & del río Buritaca-La Cumbre). In: Rangel, 1. D. OTTE. 1995. New grasshopper gene- O. (ed.), Colombia: Diversidad biótica 1, Ins- ra and species from the Dominican tituto de Ciencias Naturales, Convenio Republic (Hispaniola) (Acridoidea: Inderena-Universidad Nacional de Colom- Acrididae). Trans. Amer. Entomo!. Soco bia, Santafé de Bogotá, pp. 155-170. 121: 153-171. RANGEL,J. O., P. Lowv & M. AGUILAR.1995c. PINTO-DA-RoCHA,R. 1997. Systematic revision Marco general y alcances del estudio. In: of the Neotropical family Stygnidae Rangel, J. O. (ed.), Colombia: Diversidad (Opiliones, Laniatores, Gonyleptoidea). biótica 1, Instituto de Ciencias Naturales, Arq. Zool., Sáo Paulo 33: 163-342. Convenio Inderena-Universidad Nacional PLATNICK,N. I. 1975. A revision ofthe South de Colombia, Santafé de Bogotá, pp. 17-24. American spider genus Trachelopachys RAPOPORT,E. H. 1968. Algunos problemas (Araneae, Clubionidae). Am. Mus. Novit. biogeográficos del nuevo mundo con es- 2589: 1-25. pecial referencia a la región Neotropical. POWELL,R., R. W. HENDERSON,K. ADLER& H. In: Delamare Debouteville & E. H. Rapoport A. DUNDEE. 1996. An annotated checklist (eds.), Biologie de l' Amerique Australe, 4, of West Indian amphibians and reptiles. CNRS, Paris, pp. 55-110. In: Powell, R. & R. W. Henderson (eds.), RATCLlFFE, B. C. 1988. New species and Contributions to West lndian herpetology: distributions ofNeotropical Phileurini and A tribute to Albert Schwartz, Society for a new Phileurine from Burma (Coleoptera: the Study of Amphibians and Reptiles, Scarabaeidae: Dynastinae). Cols. Bull. 42: Ithaca, pp. 51-93. 43-55. PREGILL, G. K. 1981. An appraisal of the RATCL!FFE,B. C. & A. C. DELOYA.1992. The vicariance hypothesis of Caribbean biogeography and phylogeny of biogeography and its app1ication to West Hologymnetis (Coleoptera: Scarabaeidae:

72 Juan J. Morronc Cetoniinae) with a revision of the genus. Chirodamus Haliday sensu lato y revisión Co!s. Bu!!' 46: 161-202. de Pompilocalus gen. nov. (Hymenoptera, RArcLlFFE,B. C. & M. A.lvIE. 1998. New species Pompilidae). Rev. SocoEntomo!. Argent. 47: of Hemiphilcurus Ko1be (Coleop-tera: 79-84. Scarabaeidae: Dynastinae) from the ROSEN, D. E. 1976. A vicariance model of Dominican Republic with a key to the West Caribbean biogeography. Sysl. Zool. 24: Indian species of Hemiphileurus. Cols. 431-464. Bull. 52: 20 1-208. ROSEN,D. E. 1985. Geological hierarchies and RAUCIIENI3ERGER, M. 1988. Historical biogeographic congruence in the Caribbean. biogeography of Poeciliid fishes in the Ann. Missouri Bot. Gard 72: 636-659. Caribbean. Syst. Zool. 37: 356-365. RYAN, R. M. 1963. The biotic provinces of RIFKIND,J. 1994. New species of Mexican Central America. Acta Zoo!. Mex. 6: 1-55. Enoc!erus Gahan (Coleoptera: Cleridae). RZEDOWSKI,1. 1978. Vegetación de México. Cols. Bul!. 48: 283-292. Limusa, México D.F. RINGUELET, R. A. 1975. Zoogeografía y SAVAGE,J.M. 1982. The enigma ofthe Central ecología de los peces de aguas continen- American herpetofauna: Dispersal or tales de la Argentina y consideraciones vicariance? Ann. Missouri Bol. Gard 69: sobre las áreas ictiológicas de América del 464-547. Sur. Ecosur 2: 1-122. SCIIOLTZ,C. H. 1990. Revision ofthe Trogidae RIVAS,L. R. 1986. Comments on Briggs (1984): of South America (Coleoptera: Scara- Freshwater fisehes and biogeography of beoidea). J. Nal. Hist.24: 1391-1456. Central America and the Antilles. Syst. SCHUH,R. T. & M. D. SCHWARTZ.1985. Revision Zoo!. 35: 633-639. of the plant bug genus Rhinacloa Reuter RIVAS-MARTÍNEZ,S. & G. NAVARRO.1994. Mapa with a phylogenetic analysis (Hemiptera: biogeográfico de Suramérica. Madrid. Miridae). Bul!. Amer. Mus. Nat. Hist. 179: RIVAS-MARTÍNEZ,S. & O. TOVAR.1983. Síntesis 382-470. biogeográfica de los Andes. Collect. Bol. SCHULTZE-KRAFT,R. & R. 1. WILLlAMS.1990. (Barcelona) 14: 515-521. Una nueva especie de Centrosema (Df.;) RODRÍGUEZ,e. & c. FRAGOSO.1995. Especies Benth. (Leguminosae: Papilionoidae) del nuevas de Zap atadrilus James, 1991 Orinoco. Caldasia 16: 133-137. (Oligochaeta: Megascolecidae) registradas SCIIUSTER, J. & P. REYES-CASTILLO. 1990. en Cuba. Acta Zool. Mex., 11. s. 64: 21-33. Coleoptera, Passalidae: Ogypes Kaup, re- RODRÍGUEZ,G. 1992. Thefreshwater crabs of visión de un género mesomericano de America: Family Trichodactylidae and montaña. Acta Zool. Mex., 11. s. 40: 1-49. supplement to family Pseudotelphusidae. SEIDEL, M. E. 1996. Current status of Editions de lOrstom, Collection Faune biogeography of the West Indian turtles Tropicale 31, pp. 189. in the genus Trachemys (Emydidae). In: RODRÍGUEZ,G. & M. R. CAMPOS.1989. Cladistic Powell, R. & R. W. Henderson (eds.), relationships of fresh-water crabs of the Contributions to West Indian herpetology: tribe Strengerianini (Decapoda: A tribute to Albert Schwartz, Society for Pseudotelphusidae) from the northern An- the Study of Amphibians and Reptiles, des, with comments on their biogeography Ithaca, pp. 169-174. and descriptions of new species. J. Crust. SHANNON,R. C. 1927. Contribución a los estu- Biol. 9: 141-156. dios de las zonas biológicas de la Repúbli- ROIG-ALSINA,A. 1989. La posición sistemáti- ca Argentina. Rev. Soco Entomo!. Argent. ca de los grupos hasta ahora incluidos en 4: 1-14.

73 Toward a cladistic model Ior the Caribbean subregion SHEAR,W. A. & S. B. PECK. 1992. Centipeds monogr. no. 5, Universidad Nacional de Co- (Chilopoda) and Symphyla of the lombia, Santafé de Bogotá, pp. 173. Galapagos islands, Ecuador. Can. 1. Zoo!. TRYON,R. 1972. Endemic areas and geographic 70: 2260-2274. speciation in tropical American ferns. SIl3LEY,C. G. & B. L. MONROE.1990. Distri- Biotropica4: 121-131. bution and taxonomy ofbirds ofthe world. TUBERQUIA,D. 1997. Cuatro especies nuevas Vale University Press, New Haven & de Cyclanthaceae de Colombia. Caldasia London, 1111 pp. 19: 179-189. SITES,J. W. Jr., S. K. DAvls, T. GUERRA,J. B. V ALENTINE,B. D. 1991. Four new species of ¡VERSON& H. L. Snell. 1996. Character Phaenotheriopsis from Hispaniola and congruence and phylogenetic signal in Puerto Rico (Coleoptera: Anthribidae). molecular and morphological data sets: A Cols. Bul!. 45: 331-340. case study in the living iguanas (Squamata, V ANIN, S. A. 1986. Systematics, cladistic Iguanidae). Mol. Bio!. Evo!. 13: 1087-1105. analysis, and geographical distribution of SLATER,A. A. 1988. Zoogeography of West the tribe Erodiscini (Coleoptera, Curcu- Indian Lygaeidae (Hemiptera)./n: Liebherr, lionidae, Otidocephalinae). Revta. Bras. 1. K. (ed.), Zoogeography of Caribbean Entamo!. 30: 427-670. insects, Cornell University Press, lthaca & VAlU,R. P. 1989a. Systematics ofthe Neotro- London, pp. 38-60 .. pical Characiform genus Curimata Bosc STONEDAHL,G. M. & M. D. SCIIWARTZ.1994. (Pisces: Characiformes). Smithson. Two new species of Atractotomus from Contrib. Zool. 474: 1-63. southern Mexico (Heteroptera: Miridae: VARI,R. P. 1989b. Systematics ofthe Neotro- Phylinae).1. New York. Entamo!. Soco 102: pical Characiform genus Pseudocurimata 374-390. Fernández- V épez (Pisces: Ostariophysi). SULLlVAN, J., J. A. MARKERT & C. W. Smithson. Contrib. Zool. 490: 1-28. KILPATRICK. 1997. Phylogeography and VARI, R. P. 1991. Systematics ofthe Neotro- molecular systematics ofthe Peromyscus pical Characiform genus Steindachnerina aztecus species group (Rodentia: Muridae) Fowler (Pisces: Ostariophysi). Smithson. inferred using parsimony and likelihood. Contrib. Zool. 507: 1-118. Syst. Bio!. 46: 426-440. VARI,R. P. 1992a. Systematics ofthe Neotro- SZUMIK,C. A. 1994. Oligembia vetusta, a new pical Characiform genus Cyphocarax fossil Teratembiid (Embioptera) from Fowler (Pisces: Ostariophysi). Smithson. Dominican amber. 1. New York Entamo!. Contrib. Zool. 529: 1-137. Soc.102:67-73. VARI, R. P. 1992b. Systematics of the T AKHTAJAN,A. Floristic regions ofthe world. Neotropical Characiform genus University ofCalifornia Press, Berkeley. Curimatella Eigenmann and Eigenmann THOMAS,M. C. 1994. A new species and a new (Pisces: Ostariophysi), with summary record of Catogenus Westwood from the comments on the Curimatidae. Smithson. Dominican Republic (Coleoptera: Passan- Contrib.Zoo!.533: 1-48. dridae).1. New YorkEntomol Soco 102:476480. WATROlJS,L. E. 1982. Revision ofthe Neotropical TOLEDO,V. H. 1997. Revisión taxonómica del Archaeoglenes Broun (Coleoptera: género Lagocheirus Dejean para México Tenebrionidae). Cols.Bull.36: 135-142. y Centroamérica (Coleoptera: Ceramby- WII3MER,G. 1. 1989. Revision oftheweevil genus cidae).FoliaEntomo!. Mex. 101: 1-58. Tyloderma Say (Col.: Curculionidae) in TORRES,1. H. 1988. Podocarpaceae./n: Pinto, Mexico, Central America, South America, and P. & G. Lozano (eds.), Flora de Colombia, the West Indies. Evo!. Monogr. 11: 3-118.

74 Juan J. Morrone

WILLIAMS,E. E. 1989. 01d problems and new subgénero Littaea (Agavaceae) de la Sie- opportunities in West Indian biogeography. rra Madre Oriental, México. Acta Bol. Mex. Biogeogr. West Indies, 1989: 1-46. 32:47-52. WILSON,E. O. 1988. The biogeography ofthe ZlININO,M. 1990. Antillean Scarabaeidae (Co- West Indian ants (Hymenoptera: leoptera): Some problems ofphylogenetic Formicidae). In: Liebherr, l. K. (ed.), and geographic affinities. In: International Zoogeography of Caribbean insects, Symposium on Biogeographical Aspects Cornell University Press, Ithaca& London, of'Insularity, Atti dei Convegni Lincei 85, pp. 214-230. Accademia Nazionale dei Lincei, Rome, pp. ZAMlIDIO-RUIZ,S. & E. SÁNcHEz-MARTÍNEz. 715-727. 1995. Una nueva especie de Agave del

Appendix l. Acronyms ofthe authors in the taxonomic lists.

AA: Aranda el al., 1997; AB: Aalbu & 1966; DP: Demange & Pereira, 1985; DS: Triplehorn, 1991; AD: Andrade, 1995; AM: Desender el al., 1991; DT: Desender el al., Amorim & Pires, 1996; AN: Andrews, 1994; 1992a; DU: Desender el al., 1992b; DY: Deloya, AP: Andrews, 1998; AR: Arriaga el al., 1997; 1995; EM: Emmons, 1990; ES: Escobar, 1988; AS: Ashe& Lingafelter, 1995; AV: Alberico& FA: Frank, 1981; FB: Fernández-Alonso, 1997; Velasco, 1994; BA: Baert, 1994; BD: Bordón, FC: Fernández & Cogollo, 1992; FE: Freire, 1991; BE: Bernal, 1986; BK: Burke & Kovarik, 1993; FL: Flint, 1996; FN: Fernández el al., 1986; BO: Bordón, 1997; BR: Berry, 1982; BS: 1997; FO: Forero el al., 1983; FR: Franco & Bickel & Sinclair, 1997; BU: Bauer, 1993; BZ: Berg, 1997; FU: Funk, 1982; GB: Giesbert, 1994; Báez, 1977; CA: Clark, 1987; CB: Coscarón el GD: Gordon, 1977; GE: González-Elizondo & al., 1996; Ce: Coscarón & Coscaron-Arias, González-Elizondo, 1992; GH: Génier & 1995; CD: Camargo & Moure, 1996; CE: Casti- Howden, 1999; GI: Grimaldi, 1988; GO: llo & Reyes-Castillo, 1984; CF: CarIton, 1989; González, 1990; GN: Gordon, 1982; GR: Gordon, CG: Carranza-González, 1993; CH: Campos, 1994a; GS: Gordon, 1994b; GT: Grimaldi el al., 1997; CI: Cigliano & Lange, 1998; CK: Clark, 1993; GU: Gutiérrez, 1995; GY: Guyer& Savage, 1988; CL: Clark, 1992; CM: Campbell, 1988; 1986; HA: Herman, 1972; HB: Hedges, 1996b; CN:CampbeIl, 1991;CO:Coscarón, 1987;CP: HD: Howden, 1988; HE: Herman, 1975; HF: Cracraft & Prurn, 1988; CQ: Cohen, 1998; CR: Hernández el al., 1992a; HG: Hernández el al., Clark, 1993; CS: Castillo el al., 1988; CT: 1992b; HH: Hernández el al., 1992c; HI: Crother el al.. 1986; CU: CampbeIl & Peck, Herman, 1986; Hl: Howden, 1994a; HK: 1989; CV: Campbell & Lamar, 1989; CW: Ca- Howden, 1994b; HL: Hamilton, 1988; HM: brera & Willink, 1973; DA: Dorado & Arias, Hamilton, 1994; HN: Herrnan, 1991; HO: 1992; DB: Delgado & Blackaller-Bages, 1997; Howden, 1992; HS: Hespenheide, 1997; HT: DE: Deloya & Ratcliffe, 1988; DH: Delgado & Hamilton, 1997; HV: Harold & Vari, 1994; IS: Hernández, 1998; DL: Duellman, 1989; DN: Ivie & Sikes, 1995; lA: lameson, 1990; KE: Donoso-Barros, 1965; DO: Donoso-Barros, Kevan, 1977; KI: Kornicker & I1iffe, 1989; KL:

75 Toward a c1adistic model for the Caribbcan subregion

Kluge, 1988; KS: Kohlman & Sánchez, 1984; to-da-Rocha, 1997; PK: Peck & Kukalová- LA: Liebherr, 1986; LC: Lynch, 1996; LO: Peck, 1995; PL: Pérezel al., 1995; PO: Powell Lydeard el al., 1995; LE: Liebherr, 1988b; LF: el al., 1996; PT: Patton, 1987; RA: Lourenco & Flores, 1989; LG: Lattke & Goitía, Rauchenberger, 1988; RC: Rodríguez & Cam- 1997; LH: Lobo & Halffter, 1994; L/: Liebherr, pos, 1989; RO: Ratcliffe & Oeloya, 1992; RE: 1988d; LL: Llorente el al., 1997; LN: Lanteri, Ratcliffe & lvie, 1998; RF: Rodríguez & Frago- 1992; LO: Liebherr, 1997; LR: Lourenco, 1990; so, 1995; RH: Ramírez-García & Hernández- L Y: Lynch, 1982; LZ: Lizarralde de Grosso, Ortiz, 1994; RI: Rifkind, 1994; RJ: Roig-Alsina, 1982; MA: Maas & Maas-van de Kamer, 1988; 1989; RL: Ratcliffe, 1988; RM: Ramos, 1988; MB: Murillo, 1988; MC: McHugh, 1990; MO: RO: Rodríguez, 1992; RV: Rivas, 1986; RZ: Mora, 1984; MF: Morón & Ratcliffe, 1996; MG: Rzedowski, 1978; SA: Savage, 1982; SC: Magalhaes, C. & M. Türkay, 1996a; MI: Miller, Scholtz, 1990; SE: Seidel, 1996; SI: Sibley & 1995; ML: Morrone & Lopretto, 1996; MN: Monroe, 1990; SL: Slater, 1988; SP: Shear & Monserrat, 1997; MO: Morrone & Coscarón, Peck, 1992; SR: Schuster & Reyes-Castillo, 1996; MR: Morón, 1995; MS: Mathis, 1980; 1990; SS: Stonedahl & Schwartz, 1994; ST: MT: Magalháes & Türkay, 1996b; MU: Müller, Siteselal., 1996;SU:Sullivanetal., 1997;SV: 1973; MV: Méndez-Larios & Villaseñor, 1995; Schuh & Schwartz, 1985; SW: Schultze-Kraft NH: Navarrete-Heredia, 1997; NI: Nichols, & Williams, 1990; SZ: Szumik, 1994; TB: 1988;NW:Nowak, 1991;OB:O'Brien, 1986; Tuberquia, 1997; TH: Thornas, 1994; TL: OR: Ortiz-Oíaz, 1993; OS: Osella, 1980; OW: Toledo, 1997; TO: Torres, 1988; TR: Tryon, O'Brien & Wibmer, 1989; PA: Parenti, 1981; 1972; VA:Valentine, 1991; VI: Vari, 1989a; VN: PB: Platnick, 1975; PC: Pacheco & Patterson, Vanin, 1986;VR:Vari, 1989b;VS:Vari, 1991; 1992; PO: Pereira& Oemange, 1997; PE: Pereira VT: Vari, 1992a; WA: Watrous, 1982; WI: el al., 1997; PF: Papenfuss & Wake, 1987; PG: Wilson, 1988; WR: Wibmer, 1989; ZA: Page & Lydeard, 1994; PH: Peck, 1991; PI: Pin- Zamudio-Ruiz & Sánchez-Martinez, 1995.

Recibido: agosto 1012000 Aceptada: octubre 1012000

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