Description and Phylogenetic Relationships of Two New Species of Baripus (Coleoptera: Carabidae: Broscini) and Considerations Regarding Patterns of Speciation

ANNALS OF CARNEGIE MUSEUM

Vo l . 77, Nu m b e r 1, Pp. 211–227 30 Ju n e 2008

DESCRIPTION AND PHYLOGENETIC RELATIONSHIPS OF TWO NEW SPECIES OF BARIPUS (COLEOPTERA: CARABIDAE: BROSCINI) AND CONSIDERATIONS REGARDING PATTERNS OF SPECIATION

Se r g i o A. Ro i g -Ju ñ e n t Laboratorio de Entomología. Instituto Argentino de Investigaciones de las Zonas Áridas (IADIZA). Avenida Ruiz Leal s/n – Pq. Gral. San Martín, 5500 – Mendoza, Argentina [email protected]

Fe d e r i c o Ag r a i n * [email protected]

Ro d o l f o Ca r r a r a * [email protected]

Ei d e r Ru i z -Ma n z a n o s * [email protected]

Ma r c e l o F. To g n e l l i * [email protected]

Abstract

Baripus Dejean is a genus of the family Carabidae restricted to southern South America. Three subgenera are recognized within the genus; one of them, Cardiophthalmus Curtis, is endemic to the Patagonian Steppe. Together with the members of the genus Cnemalobus Guérin-Ménéville, these beetles are the largest Carabidae of the Patagonian Steppe. New studies in the northern region of Patagonia, the Payunia, reveal the presence of two new species, Baripus (Cardiophthalmus) nevado, new species, and Baripus (Cardiophthalmus) precordillera, new species, restricted to isolated montane habitats. In this paper, we provide morphological descriptions of the new taxa with illustrations of male and female genitalia. We also conducted a phylogenetic analysis including all known species of Baripus. The cladistic analysis showed that B. (C.) mendozensis, B. (C.) nevado and B. (C) precordillera constitutes a monophyletic group of species. These three species are allopatric, but B. (C.) mendozensis and B. (C.) nevado ranges are in close proximity. The distribution pattern, together with the phylogenetic pattern, suggests a possible peripatric pattern of speciation. We used a predictive model of species distribution to establish the present, past, and future distribution of B. (C.) mendozensis which is the most widely distributed species. Based on these analyses, we speculate that the current pattern of speciation may be a result of climatic changes during the Pleistocene.

Ke y Wo r d s : Baripus, Broscini, Carabidae, Cardiophthalmus, Coleoptera, new species, pattern of speciation, phylogeny, vicariance.

RESUMEN

Baripus Dejean es un género de la familia Carabidae del Sur de Sudamérica. Se reconocen tres subgéneros dentro del género; uno de ellos, Cardiophthalmus Curtis, es endémico de la estepa de Patagonica. Junto con los miembros de Cnemalobus Guérin Ménéville, son los géneros de Carabidae de mayor tamaño de la estepa patagónica. Nuevos estudios en la región norte de Patagonia, Payunia, revelan la presencia de dos nuevas especies Baripus (C.) nevado, n. sp., y B. (C.) precordillera, n. sp., restringidas a ambientes montanos aislados. Se suministran descripciones morfológicas con ilustraciones, incluyendo genitalia femenina y masculina, para los nuevos taxa. Los análisis filogenéticos fueron desarrollados utilizando todas las especies conocidas de Baripus. El análisis cladístico muestra que B. (C.) mendozensis, B. (C.) nevado and B. (C) precordillera constituyen un grupo monofiletico. Estas tres especies son alopátricas, pero en B. (C.) mendozensis y B. (C.) nevado los límites de sus áreas de dis- tribución son muy cercanos. El patrón de distribución, junto con el filogenético, muestra un posible patrón de especiación peripátrica. Se utilizó un modelo predictivo de distribución de especies para establecer la distribución presente, pasada y futura de B. (C.) mendozensis, la cual es la especie mas ampliamente distribuida. Sobre la base de estos análisis, especulamos que los cambios climáticos durante el Pleistoceno podrían haber causado el patrón de especiación actual.

Pa l a b r as Cl a v e : Baripus, Broscini, Carabidae, Cardiophthalmus, Coleoptera, filogenia, nueva especie, patrones de especiación, vicariancia.

*Same address as senior author. 212 An n a l s o f Ca r n e g i e Mu s e u m Vo l . 77

INTRODUCTION anatomical structures. To study the male genitalia, the aedeagus was removed through the pygidium and cleared in Baripus Dejean is a South American genus of the tribe boiling 10% KOH for ten minutes. The internal sac was Broscini, comprising a monophyletic group comprised of everted by injecting 1:1 glycerin with a hypodermic sy- species classified in three subgenera (Roig-Juñent 1992b, ringe. To study the female genitalia, membranes and mus- 2000). The systematics of the genus were recently stud- cles were removed. All the illustrations were made using a ied by Roig-Juñent and Cicchino (1989) and Roig-Juñent camera lucida attached to a Nikon stereomicroscope. Pic- (1992a, 1992b). The phylogenetic and biogeographical tures of male protarsomeres were taken with a Power Shot hypotheses were analyzed by Roig-Juñent (1995, 2000) S50 digital camera adapted to a Leica MS5 stereoscopic to investigate vicariant events in the history of Baripus. microscope. These pictures were taken at different focal The three subgenera exhibit an allopatric distribution and lengths, and then a single image was obtained by using inhabit different types of grassland. The most diverse sub- Helicon Focus v3.20 software, available at www.helicon- genus, Cardiophthalmus Curtis (1839), occurs exclusively focus.com. The terms used for the aedeagus follow Jeannel in the Patagonian Steppe, a lowland habitat that extends (1955), and the terms used for the female genitalia follow from the Atlantic Coast to the western slope of the Andes Deuve (1993) and Liebherr and Will (1998). Mountains. After the systematic revisions (Roig-Juñent and Cicchino 1989; Roig-Juñent 1992a, 1992b), we collected new specimens of the known species of the subge- Cladistic Analysis nus Cardiophthalmus and obtained new distributional data. Characters.—The 55 characters and their states used for We also discovered two new species of Cardiophthalmus, the analysis are shown in Table I. The distribution of the one in the northern region of Patagonia, and the other in states among the terminal taxa are in the data matrix (Table the southern region of the Precordillera Formation. These 2), and the apomorphic states are shown in the obtained new records allowed us to analyze accurately the distribu- cladograms (Fig. 8). When a character had more than two tion pattern of Cardiophthalmus species. We then used the states, the states were organized in the matrix with most phylogenetic relationships of the species to determine their similar states adjacent, and most divergent states at op- geographical pattern of speciation. posing ends of the series (i.e., characters 8, 17, 36, and The objectives of this study were to describe two new 53). Character 53 was analyzed as an ordered multistate species of Baripus, to develop a hypothesis of the phyloge- character; all others were analyzed as unordered. Polymor- netic relationships of the new taxa, to describe the habitats, phic characters are represented by letters in the data matrix and to analyze the possible biogeographic hypotheses that (Table 2). led to the current distribution pattern of the genus. We in- ferred a phylogeny for the lineage, and used it to establish Taxa.—All species of the genus Baripus were included in possible patterns of speciation by comparing pairs of sister the analysis, 24 known species and the new taxa Baripus species and sister clades. We also used predictive models (C.) nevado, new species, and Baripus (C.) precordillera, with an exemplar species to determine likely events influ- new species. The cladograms obtained were rooted follow- encing past and future distributions. ing the Parsimonius Outgroup Analysis Method proposed by Farris (1982) and suggested by Nixon and Carpenter MATERIALS AND METHODS (1993). This method involves adding an external group to the matrix, analyzing it without forcing any type of rela- Specimens and Taxonomic Material.—Most of the spec- tionship, and rooting between the external and the internal imens we examined belong to the Instituto Argentino de group. As external groups, we chose the genera Nothobro- Investigaciones de las Zonas Áridas (IADIZA), Mendoza, scus Roig-Juñent and Ball (Nothobroscus chilensis Roig- Argentina. We also used specimens from Museo Argentino Juñent and Ball), Creobius Guérin Ménéville (Creobius de Ciencias Naturales “Bernardino Rivadavia” (MACN), eydouxii (Guérin Ménéville)), Cascellius Curtis (Cascel- Museo de La Plata (MLP), American Museum of Natural lius gravesii Curtis), and Bembidiomorphum Champion History (AMNH), and Smithsonian Institution, National (Bembidiomorphum convexum Champion). These four Museum of Natural History (USNM). We studied a total of genera belong to different subtribes of the tribe Broscini 374 specimens of Baripus (Cardiophthalmus) mendozensis (Roig-Juñent 2000): Nothobroscus to Nothobroscina, Cre- Roig-Juñent and the new species to analyze their morphol- obius and Cascellius to Creobiina, and Bembidiomorphum ogy. We also studied a series of 237 specimens of other (=Microbarypus Roig-Juñent) to Barypina. species of Cardiophthalmus held at different museums and institutions to establish geographical distribution patterns. Data Analyses.—The data matrix was analyzed with the program TNT (Goloboff et al. 2003) using the implicit enu- Dissection Methods.—The comparisons among the dif- meration search option, which finds the most parsimonious ferent species were based on characteristics of the exter- trees by means of an exact solution. Branch support was nal morphology and male and female genitalia. Speci- measured using two resampling methods with 100 repli- mens were soaked in NH3 for ten minutes to relax their cates: Jackknifing (Lanyon 1985; Farris et al. 1996), p=36 2008 Ro i g -Ju ñ e n t e t a l .—Tw o Ne w Sp e c i e s o f Ba r i p u s 213

Ta b l e 1. Characters and character states used in the phylogenetic analysis of Baripus.

Coloration 1.—Coloration: nonmetallic (0); metallic (1).

Head 2.—Setae of submentum: four (0); two (1); six (2). 3.—Tooth of mentum: apex shallowly emarginate (0); deeply emarginate, with lateral projections (1); rounded, not emarginate (2). 4.—Longitudinal medial carina of glossal sclerite: absent (0); small (1); pronounced (2). 5.—Number of glossal sclerite setae: two (0); four (1). 6.—Maxillary palpomeres: slender (1); broad (2). 7.—Number of basal stipital setae: two (0); three (1). 8.—Antennomeres 4–10: one times longer than broad (0); two times longer than broad (1); three times longer than broad (2). 9.—Row of posterolateral setae antennomeres 2–5: absent (0); present (1). 10.—Frontal grooves: absent (0); shallow (1); deep, thin (2); deep broad (3). 11.—Eyes: rounded (0); with emargination in front of antennal insertion (1).

Prothorax 12.—Setae of pronotum: one (0); two (1); three (2); four (3); five (4); six (5). 13.—Prosternal setae: absent (0); present (1). 14.—Setae of protrochanters: two (0); one (1). 15.—Protibiae: not prolonged externally to a point at apex (0); prolonged externally (1). 16.—First protarsomere male: expanded on inner side (0); not expanded (1); quadrangular (2). 17.—Squamosetae of male protarsomeres: absent (0); restricted to a small patch or few setae on the apical margin (1); apical (from the middle) (2); over entire surface (3); just three or four setae on the apical margin (4). 18.—Protarsomeres female: not expanded on inner side (0); expanded on inner side (1).

Pterothorax 19.—Metepimeron: transverse and slender (0); transverse and broad (1); subquadrangular (2). 20.—Meso- and metatarsomeres: broad (0); slender (1). 21.—Male mesotarsomeres 1–2: without adhesive setae (0); with adhesive setae (1). 22.—Lateral row of setae of mesotibiae: absent (0); present (1). 23.—Setae of metafemora: in a row (0); not aligned (1). 24.—Elytral epipleura: slender (0); broad (1). 25.—Parascutellar strioles: isolated from the apical portion of stria 1 (0); joined to the apical portion of stria 1 (1). 26.—Setae of interval 3, 5, and 7: absent (0); present (1). 27.—Setae of interval 9: 3–7 (0); 8–17 (1); 19–29 (2); 30–51 (3).

Abdomen 28.—Abdominal sterna: smooth (0); lateral basal sulcus and a shallow fovea in each side (1); with a complete basal sulcus (2). 29.—Anal setae: separate centrally (0); not separate (1). 30.—Anal setae: Three to eight (0); 10–20 setae (1); 21–40 setae (2). 31.—Apical groove of abdominal sterna: absent (0); present (1).

Male Genitalia 32.—Left paramere: setose (0); asetose (1). 33.—Basal keel of median lobe: absent (0); present (1). 34.—Basal orifice of median lobe: closed dorsally (0); open (1). 35.—Median lobe, basal expansions: absent (0); present (1). 36.—Dorsal surface of median lobe: completely sclerotized (0); partially sclerotized (1); not sclerotized (2). 37.—Lateral apical groove of median lobe: absent (0); short (1); long and straight (2); long and curved (3). 38.—Lateral grooves of median lobe: absent (0); present curved (1). 39.—Ventral groove of median lobe: absent (0); slender (1); broad (2). 40.—Ventral doubling of median lobe: absent (0); present (1). 41.—Tooth in the medial region of basal surface of median lobe: absent (0); present (1). 42.—Ventral doubling of ostium of median lobe: absent (0); present (1). 43.—Ostium of median lobe: long and broad dorsolateral (0); broad, left lateral (1); small dorsoapical (2); small, left lateral (3). 44.—Apex of median lobe: wide, subquadrate (0); wide, rounded, downward (1); long and sharpened (2); spatulate (3). 45.—Tooth of internal sac: absent (0); conical, broad (1); spatulate (2); conical slender (3); short (4); conical, very long (5). 46.—Sclerite alpha of internal sac: absent (0); present, formed by unfused spiculae (1); formed by fused spiculae (2). 47.—Apical plate of internal sac: present (0); absent (1). 48.—Sclerites X and Y of internal sac: present (0); absent (1). 49.—Internal sac eversion: from the dorsum (0); from the left (1).

Female Genitalic Tract 50.—Median oviduct union on bursa copulatrix: apical (0); basal (1). 51.—Spermathecal union on bursa copulatrix: sessile (0); with median oviduct (1). 52.—Accessory gland of bursa copulatrix: present (0); absent (1). 53.—Pocket of apex of bursa copulatrix: absent (0); small, with a portion sclerotized (1); long, completely sclerotized (2). 54.—Gonopod VIII: long and slender (0); short (1); long and broad (2); absent (3). 55.—Helminthoid sclerite: absent (0); present (1). 214 An n a l s o f Ca r n e g i e Mu s e u m Vo l . 77

Ta b l e 2. Data matrix of species of Baripus. Note: polymorphic entries: [01] = a, [12] = b, [23] = c, and missing data = “?”.

1 2 3 4 5 Nothobroscus 0112000001000000000000001010000000000000000?00000000000 Bembidiomorphum 1201001000101000200010000010000010000000002000110011001 Creobius 1120100100040002200010001011000101122000000000110010031 Cascellius 1120100100010002200010001001001101122000000000110010031 B. parallelus 0112001010131001000000100002001110010000000000110011011 B. bonvouloirii 0112001010131001000000100002001110010000000000110011011 B. rivalis 1012001011111010111000100002001110001010001011111011011 B. pulchellus 1012001012121011011100100012001110001000001821111011021 B. speciosus 1012011013151011001000100012001110001000001421111011021 B. aequicostis 10120110131510110?1000100012001110001000001021111?????? B. comechingonensis 1012001012131011001000100012001110001000001432111011021 B. calchaquensis 1012001012121011001000100012001110001000001432111011021 B. longitarsis 0112001210111110012101110132121110002020001400111111001 B. schajovskoii 01120012101211100121011101321b1110000021001010111111001 B. gentilii 01120012101211100121011101321b1110002021001010111111001 B. mendozensis 01120012101211103121011100c3121110002021101300111111001 B. precordillera 0112001210121110412101110022121110002021101300111111001 B. nevado 0112001210121110012101110022121110002021101000111111001 B. flaccus 0112001110111110012101110032111110002000001600111111001 B. painensis 0112001110111110112101110032111110003110011010111111001 B. deplanatus 0112001110111110112101110032111110003110011510111?????? B. sulcatipenis 0112001110111110112101110032111110003100011010111111001 B. neuquensis 0112001110121112212101110022001110000000003710111011001 B. giaii 01120011101211122121011100b2001110000000003250111011201 B. dentipenis 0112001110121112212101110032001110000000003250111011201 B. chubutensis 01120011101211122121011100b2001110000000003140111011101 B. minus 0112001110121112212101110022001110000000003140111011101 B. clivinoides 01120011111311101121011100b20a1110000000003011111011001 2008 Ro i g -Ju ñ e n t e t a l .—Tw o Ne w Sp e c i e s o f Ba r i p u s 215

(p= removal probability) and Bootstrapping (Felsenstein 1985). Support numbers were given as relative values. Predictive Models of Species Distribution. In general, species distribution models are used to predict species potential distribution by relating the known locations of species collection to a set of environmental variables that, presumably, reflect the ecological niche of the species (Guisan and Thuillier 2005). These models produce spatial predictions of habitat suitability that indicate where the target species may occur. The modeled niches can be projected under different environmental scenarios, thus producing habitat suitability maps for past or future climatic conditions (Graham et al. 2004; Yesson and Cul- ham 2006). In this study, the presently known locations of Baripus (C.) mendozensis were georeferenced, mapped, and used to model its potential distribution. Species distribution models were run using present, past (21,000 years B.P.), and future (year 2050) climate data. We used the program MaxEnt (Phillips et al. 2006), combined with 19 bioclimatic variables obtained from the WorldClim dataset (Hijmans et al. 2005) to model the distribution of B. (C.) mendozensis. The resolution of the environmental layers used was, approximately, 4.6 x 4.6 km.

SYSTEMATIC ENTOMOLOGY Fig. 1.—Baripus (Cardiophthalmus) nevado, habitus. Order Coleoptera Linnaeus, 1758 Family Carabidae Latreille, 1802 lateral and 0.5 as long as length of median lobe (Fig. 2A); Tribe Broscini Hope, 1838 internal sac without tooth; spermatheca and oviduct joining Genus Baripus Dejean, 1828 together at base of oviduct (Fig. 2F). Subgenus Cardiophthalmus Curtis, 1839 Description.—Habitus as in Fig. 1. Length: 15.45–18.35 mm (x=16.92). Baripus nevado Roig-Juñent, new species Coloration.—Black, with mouthparts, antennae, and legs testaceous dark reddish. ( Figs. 1–4) Head.—Labrum short, transverse, bilobate at anterior margin. Clypeus with two subparallel lateral sulci slightly developed, each with one seta. Type Material.—Holotype: Argentina. Mendoza. Malargüe: Cerro El One posterior supraorbital seta. Eyes with anterior emargination near Nevado, 35°37’40”S, 68°32’40”W, 2361 m, 1 male (IADIZA). Paratypes: antennal insertion. Antennomeres three times as long as wide. labeled as holotype 1 male, 2 females (IADIZA); El Nevado, arbustal de Pronotum.—Wider than long (width/length= 1.19), maximum width Adesmia, 35°36’06”S, 68°30‘51”W, 2925 m, 7 January 2006, G. Debandi at apical third; dorsal surface smooth. Lateral margins narrow, curved, Collector, 57 specimens (AMNH, IADIZA, MACN, MLP, USNM); El usually with three setae on each side, two setae on apical third and one Nevado, ecotono entre estepa y altoandina, 35°36’57”S, 68°31’43”W, 2590 on basal third, some specimens with only apical two and other specimens m, 7 January 2006, G. Debandi Collector, 1 male (IADIZA); El Nevado, with four setae. Central longitudinal sulcus slightly developed; posterior 35°36’9”S, 68°32’47”W, 2935 m, February 2005, G. Debandi Collector, transverse foveae slightly impressed. Posterior angles rounded. 1 male, 4 females (IADIZA); El Nevado, 35°36’11”S, 68°30‘49”W, Elytra.—Humeral angles rounded; epipleura decreasing in width from 2908 m, December 2004, G. Debandi Collector, 12 males (IADIZA); El base to apex; striae weakly impressed, more visible at apex. Setae only in Nevado, 35°36’35”S, 68°31’41“W, 2662 m, December 2004, G. Debandi ninth interval, 28–38 setae. Collector, 1 male, 3 females (IADIZA). Legs.—Profemora with three or four teeth ventrally. Protrochanters with one seta present. Protarsomere 1 four times as long as basal width; first and Specific Epithet.—The name of the new species is related second protarsomeres of males without adhesive setae; meso- and metatar- to the mountain where it was collected, El Nevado Moun- someres long (length all metatarsomeres/ length elytra= 0.53–0.57). Males tain in Mendoza Province, Argentina, and is to be treated as mesotibiae with apical brush of setae. Abdominal Sterna.—Sterna V–VII with basal sulcus; sterna IV–VI with a noun in apposition. 1–2 setae; sternum VII with 30–38 apical setae. Male Genitalia (Figs. 2A–E) .—In addition to diagnosis, median lobe Diagnosis.—Lateral margin of pronotum narrow, with three wide (Figs. 2A–B), recurved from base to apex (Fig. 2E), with apex short setae on each side (Fig. 1); elytra with striae indistinct, with- and quadrangular (Figs. 2A–B) and orifice of median lobe 0.5 times length out setae in intervals 3, 5, 7; males protarsomeres four times of median lobe, opening on left side of median lobe. Orifice of median lobe without spiculae and consequently without ligula. Internal sac 0.5 times as as long as wide, not expanded at apex, without adhesive long as length of median lobe, without tooth (Fig. 2A), but with apical group setae; median lobe of aedeagus with a ventral groove and of free spiculae. Left paramere broad, with rounded apex and setae on apical associated ventral tooth or mucro, orifice of median lobe third (Fig. 2C); right paramere with slender apex, glabrous (Fig. 2D). 216 An n a l s o f Ca r n e g i e Mu s e u m Vo l . 77

is an orogenic mountain range that extends from north to south from latitude 35ºS to 36°S and is parallel (200 km to the east) to the Andean Mountains. The Sierras del Nevado Mountains increase in height from north to south, reaching their maximum altitude at El Nevado Mountain (3833 m). The north-south orientation of this mountain produces a marked difference in rainfall between eastern and western slopes. Baripus (C.) nevado n. sp. was found in two types of vegetation that occur at different altitudes. The first type, located at the base of the mountain, is a shrub steppe on sandy and basaltic soils with Neosparton aphyllum Gilles and Hook. (Verbenaceae) and Sporobolus rigens (Trinius) Desvaux (Gramineae). The second type, located in high volcanic plateaus, is characterized by the presence of Adesmia pinnifolia Gilles ex Hook. and Arn. (Fabaceae) and Anarthrophyllum rigidum Hieronymus (Leguminosae).

Other Material Examined.—Argentina. Mendoza. Malargüe: El Nevado, ecotono entre estepa y provincia biogeográfica altoandina, 35°36’57”S, 68°31’43”W, 2590 m, 7 January 2006, G. Debandi Collector, 2 specimens (IADIZA); El Nevado, 35°36’9”S, 68°32’47”W, 2935 m, February 2005, G. Debandi Collector, 1 male, 3 females (IADIZA); El Nevado, 35°37’44”S, 68°32’52”W, 2333 m, 19 February 2005, S. Claver Collector, 1 female (IADIZA); El Nevado, 35°37’40”S, 68°32’39”W, 2361 m, December 2004, G. Debandi Collector, 3 specimens (IADIZA); El Nevado, 35°36’11”S, 68°30‘49“W, 2908 m, December 2004, G. Debandi Collector, 72 specimens (IADIZA); El Nevado, 35°36’35“S, 68°31’41“W, 2662 m, December 2004, G. Debandi Collector, 11 specimens (IADIZA); El Nevado, 35°36’47”S, 68°31’39”W, 2607 m,

Fig. 2.—Genitalia of Baripus (Cardiophthalmus) nevado. A, aedeagus, left lateral view (M=Ventral mucro); B, aedeagus, right lateral view; C, aedeagus, left paramere; D, aedeagus, right paramere; E, aedeagus, lateral view; F, female genitalia.

Female Genitalia (Fig. 2F) .—Bursa copulatrix without sclerotized ventral side. Spermatheca long, slightly curved with narrow apex; joining the base of bursa copulatrix along with the oviduct. Stylomeres short and rounded, with one nematiform seta.

Comparative Notes.—Baripus (C.) nevado shares several characters with B. (C.) mendozensis, but there is one character, the presence of the ventral mucro on the median lobe, that is exclusively shared with this species. The most conspicuous difference between the external morphology of B. (C.) nevado and B. (C.) mendozensis is the presence of adhesive setae in male protarsomeres 1 and 2 in B. (C.) mendozensis (Fig. 5D) and the absence in B. (C.) nevado. Differences in the genitalia between both species are in the shape of the median lobe and in the presence of a ligula at the orifice of median lobe in B. (C.) mendozensis (Fig. 5A–C) and the absence in B. (C.) nevado (Figs. 2A–B).

Distribution and Habitat.—The present distribution of B. (C.) nevado is restricted to El Nevado Mountain (Fig. 4). The specimens were collected at different elevations, from 2000 m altitude at the base of the mountain to 2900 m altitude (Figs. 3A–B). The region where the species was found corresponds to the northern area of the Patagonia Biogeographic Province, Fig. 3. El Nevado Mountain in Mendoza Province, Argentina. A and B, more specifically, in the Sierras del Nevado Mountains. Sierras del Nevado two perspectives. 2008 Ro i g -Ju ñ e n t e t a l .—Tw o Ne w Sp e c i e s o f Ba r i p u s 217

February 2005, G. Debandi Collector, 9 specimens (IADIZA); El Nevado, 35º37’48”S, 68°32’14“W, 2356 m, February 2006, R. Carrara Collector, 1 specimen (IADIZA); El Nevado, 35°37’41”S, 68°32’36”W, 2354 m, December 2004, G. Debandi Collector, 36 specimens (IADIZA); El Nevado, 35°37’40”S, 68°32’39”W, 2361 m, December 2004, G. Debandi Collector, 4 males (IADIZA); 7.5 km desde entrada a El Nevado Ruta 160, 35°37’29”S, 68°36’05”W, 2001 m, 25 February 2006, Carrara & Agrain Collectors, 1 male (IADIZA).

Baripus precordillera Roig-Juñent, new species (Figs. 4, 6A–D)

Type Material.—Holotype: Argentina. Mendoza: Las Heras: Paramillos, 32°30’02“S, 69°03’60”W, February 2005, G. Debandi Collector, 1 male (IADIZA). Paratypes: labeled as holotype, 1 male, 2 females (IADIZA); Paramillos, 32°30’02”S, 69°03’05“W, February 2004, 1 female (IADIZA); Paramillos, 32°30‘03”S, 69°04‘03“W, December 2004, G. Debandi Collector, 3 males, 3 females (IADIZA); Fig. 5.—Baripus (Cardiophthalmus) mendozensis. A, median lobe of 3 km Este Paramillos de Uspallata, 2830 m, 1 November 2001, S. Roig aedeagus, left lateral view; B, aedeagus, apex of median lobe; C, median Collector, 2 males (IADIZA); Paramillos, 32°30’02”S, 69°03’60”W, lobe of aedeagus, ventral view; D, male protarsus. February 2005, G. Debandi Collector, 1 male (IADIZA); Paramillos, 32°30‘03”S, 69°04‘00”W, December 2004, G. Debandi Collector, 1 female (IADIZA); Paramillos, 32°30’29”S, 69°04’00”W, February 2005, 69°09’01”W, 2904 m, 29 December 2005 – 7 January 2006, Claver-Ferrer G. Debandi Collector, 2 females (IADIZA); Paramillos, 32°30’02”S, Collector, 1 female (IADIZA); Pampa Canota, 32°40’28“S, 69°09’03”W, 69°03’60”W, December 2004, G. Debandi Collector, 4 males (IADIZA); 2897 m, 29 December 2005 – 7 January 2006, Claver-Ferrer Collector, Paramillos, 32°30’03”S, 69°04’03” W, February 2005, S. Roig Collector, 1 male, 3 females (IADIZA); Pampa Canota, 32°40’25”S, 69° 09’04”W, 1 female (IADIZA); Paramillos, 32°30’00”S, 69°03’60”W, December 29 December 2005 – 7 January 2006, Claver-Ferrer Collector, 3 females 2004, S. Roig Collector, 1 male (IADIZA); Pampa Canota 32°40’25”S, (IADIZA); Pampa Canota 32°40‘25”S, 69°09’00”W, 2896 m, 29 December 2005 – 7 January 2006, Claver-Ferrer Collector, 2 females (IADIZA); Pampa Canota, 32°40’26”S, 69°08’59”W, 2896 m, 29 December 2005 – 7 January 2006, Claver-Ferrer Collector, 2 females (IADIZA); Pampa Canota, 32° 38.563´ S, 69° 08.202´ W, 2976 m, 15 February 2006 – 25 February 2006, Lagos-Flores Collector, 1 male, 1 female (IADIZA); Pampa Canota, 32°38’36“S, 69°08’04”W, 2973 m, 15 February 2006 – 25 February 2006, Lagos-Flores Collector, 1 female (IADIZA); Paramillos, 32°29’08“S, 69°05’21”W, 2857 m, 28 December 2005 – 6 January 2006, Sallenave-Scollo Collector, 5 males, 2 females (IADIZA); Paramillos, 32°29’09“S, 69°05’20”W, 2853 m, 28 December 2005 – 6 January 2006, Sallenave-Scollo Collector, 3 females (IADIZA); Paramillos, 32°29’13“S, 69°05’20”W, 2852 m, 28 December 2005 – 6 January 2006, Sallenave-Scollo Collector, 7 males, 2 females (IADIZA); Paramillos, 32°29’12”S, 69°05’19“W, 2875 m, 28 December 2005 – 6 January 2006, Sallenave-Scollo Collector, 1 female (IADIZA); Paramillos, 32°29’12”S, 69°05’20”W, 2849 m, 28 December 2005 – 6 January 2006, Sallenave-Scollo Collector, 2 males (IADIZA); Paramillos, 32°29’07”S, 69°05‘22”W, 2872 m, 28 December 2005 – 6 January 2006, Sallenave-Scollo Collector, 1 male, 4 females (IADIZA); Paramillos, 32°29’10”S, 69°05’19“W, 2855 m, 28 December 2005 – 6 January 2006, Sallenave-Scollo Collector, 9 males, 10 females (IADIZA). Specific Epithet.—The name of the species is related to the mountain range where the specimens were collected, the Precordillera , and is to be treated as a noun in apposition. Diagnosis.—Elytral interval 9 with 23–28 setae; elytra without striae; last abdominal sternum with 16–27 apical setae; mesotibiae with 15–20 setae; male with few (three or four) squamosetae on protarsomeres 1 and 2 (Fig. 6A); median lobe of male aedeagus curved, with ventral fossa and associated mucro, mucro with rounded apex (Figs. 6B–C); apex of median lobe spatulate (Fig. 6D); spermatheca and oviduct joined together at base of bursa copulatrix.

Fig. 4.—Baripus (Cardiophthalmus) nevado and Baripus Description.—Habitus very similar to Baripus (C.) mendozensis. Length: (Cardiophthalmus) mendozensis, distribution map. 18.57–21.48 mm (x = 19.95 mm). 218 An n a l s o f Ca r n e g i e Mu s e u m Vo l . 77

long, and runs parallel to the Andean Mountains, but is separated by about 50–100 km (Roig-Juñent et al. 2003). There are three Biogeographic Provinces in the Precordillera: the Puna, the Monte, and the Prepuna (Roig and Martínez Carretero 1998). Baripus (C.) precordillera is found in the Puna habitat, characterized for having a cold and dry climate (Roig and Martínez Carretero 1998, Roig-Juñent et al. 2003). The Puna is located between 2700 and 3000 m altitude (Figs. 7A–B), and its vegetation consists mainly of grasslands and arid shrub steppes. The steppes have small shrubs between 40–150 cm tall that are interspersed with bare soil that receives high solar irradiation. The characteristic plants in this area are Lycium fuscum Miers (Solanaceae), Ephedra mutiflora Phil. Ex Stapf (Ephedraceae), Verbena diversifolia Kuntze (Verbenaceae), Baccharis thymifolia Hook., and Arn. (Asteraceae), B. polifolia Griseb (Asteraceae) (Roig 1972). Fig. 6.—Baripus (Cardiophthalmus) precordillera. A, protarsus; B, median lobe of aedeagus, left lateral view; C, median lobe of aedeagus, Material Examined.—Holotype, Argentina, Mendoza, Malargüe: ventral view; D, apex of median lobe of aedeagus. Fortín Malargüe, Mendoza, January 10 1974, A. Roig Collector, male (IADIZA). Paratypes: a escasos kilómetros del Mollar, Mendoza, Coloration.—Black, mouth parts and some regions of legs and anten- Malargüe, January 23 1979, S. Roig Collector, 1 male (IADIZA); Agua nae dark reddish. Escondida, January 29 1979, S. Roig Collector, 1 female (IADIZA); Ruta Head.—Labrum short, transverse, anterior margins straight with small 180 El Nevado, 35°36’08”S, 68°40’16”W, February 2006, R. Carrara emargination at middle, lateral margins subparallel. Clypeus not emargin- Collector, 1 male, 1 female (IADIZA). San Carlos: Arroyo Cruz de ate, with two lateral sulci, each with one seta. Piedra, 34°14’47”S, 69°22’51”W, 2333 m, G. Debandi Collector, 1 male, Pronotum.—As wide as long (width/ length: 1.04), maximum width 1 female (IADIZA); Piedra Pomes, 34°12’28“S, 69°08’00“W, 1511 m, at middle. Lateral margins narrow, curved, with three setae on each side, 25 February 2005, R. Carrara Collector, 2 males, 9 females (IADIZA); two at anterior half and one at posterior third. Posterior foveae impressed. Arroyo Cruz de Piedra, 34°14‘46”S, 69°22‘51”W, 2324 m, 18 February Posterior angles rounded, inconspicuous. 2005, G. Debandi Collector, 6 males, 4 females (IADIZA); Refugio Elytra.—Shape ellipsoid, without developed striae, surface smooth. Alvarado, 34°14’38”S, 69°22’42“W, 2296 m, 27 December 2005 – 5 Humeral angles rounded. Setae only on interval 9, 23–28 setae present January 2006, Roig-Debandi Collector, 1 male, 1 female (IADIZA); (X= 24). Refugio Alvarado, 34°14‘46”S, 69°22‘51”W, 2329 m, 8 February 2005 – Legs.—Profemora with ventral teeth. Protrochanters with one seta. 18 February 2005, Claver-Tognelli Collector, 1 male (IADIZA); Refugio Male protarsomeres 1 and 2 with 3–4 adhesive setae on apical side (Fig. Alvarado, 34°14‘46”S, 69°22‘50”W, 2327 m, 8 February 2005 – 18 6A). Meso- and metatarsomeres long (length all metatarsomeres/length February 2005, Claver-Tognelli Collector, 7 males (IADIZA); Refugio elytra=0.48). Male mesotibiae with lateral brush consisting of 15–20 Alvarado, 34°14‘47”S, 69°22‘51”W, 2333 m, 8 February 2005 – 18 (X=15) setae. February 2005, Claver-Tognelli Collector, 2 female (IADIZA); Refugio Abdominal Sterna.—Sterna V–VII with basal sulcus; Sterna IV–VI Alvarado, 34°14‘46”S, 69°22‘51”W, 2328 m, 8 February 2005 – 18 with one or two apical setae; VII with 16–27 setae (X= 16) on apical February 2005, Claver-Tognelli Collector, 2 males, 6 females (IADIZA); margin. Refugio Alvarado, 34°14‘47”S, 69°22‘51”W, 2332 m, 8 February Male Genitalia.—(Figs. 6B–D) Median lobe curved, with apex 2005 – 18 February 2005, Claver-Tognelli Collector, 7 males, 3 females enlarged and widened, spatulate (Fig. 6D), orifice of median lobe latero- (IADIZA); Refugio Alvarado, 34°14‘46”S, 69°22‘51”W, 2327 m, 8 dorsal, with a group of spiculae at base that constitutes the ligula (Fig. February 2005 – 18 February 2005, Claver-Tognelli Collector, 5 males, 4 6A). Internal sac without tooth, with small spicules only. females (IADIZA). San Rafael: 1.5 km de Agua del Toro, 18 December 1998, Flores-Roig Collector, 1 male (IADIZA); Dique Agua del Toro, 18 Comparative Notes.—B. (C.) precordillera and B. (C.) mendozensis dif- December 1998, Flores-Roig Collector, 1 male (IADIZA). fer in their external morphology by the number of adhesive setae of male protarsomeres (Compare Figs. 5D and 6A) and the number of elytral and VII sternal setae. The male genitalia between these species are slightly dif- Key to species of the subgenus ferent (compare Figs. 5A–C with Figs. 6B–D). These differences are the Baripus (Cardiophthalmus) degree of curvature of the median lobe and the shape of the apex of median lobe and of the ventral mucro. The most important differences between 1. Antennomeres 5–10 three times as long as wide; the two species are explained in Table 3. sternum VII (last visible sternum) with setae Distribution and Habitat Characterization.—B. (C.) precordillera almost covering external margin 2 occurs in the northwestern region of the Mendoza Province (Fig. 4). Most of the specimens were collected at high altitude, between 2849 and 1’. Antennomeres 5–10 two times as long as wide; 2976 m (Figs. 7A–B) . However, some specimens were labeled as being sternum VII with setae on apical half or just with collected in Uspallata, which is located at about 1900 m altitude and cor- few setae on apex 7 responds to a completely different habitat from that of the high altitude. Because Uspallata is close to where the other specimens were collected, 2 (1). Three setae on each side of pronotum; median lobe we assumed that the material labeled as “Uspallata” was incorrectly with wide ventral fovea, not divided 3 labeled. We had three reasons for this assumption. First, the habitats of both areas are completely different: one is high altitude grassland and the 2’. Two setae on each side of pronotum; median lobe other a desert scrub area. Second, since 1989, there have been several with ventral fovea divided by a small carina at field collecting trips, and the material of the new species has always been middle B. (C.) longitarsis (Waterhouse,1841) collected at high altitudes and never in the lowland area. Third, during a exhaustive two-year ecological study in these two areas carried out by a member of our laboratory, this species was only found at high altitude. 3 (2). Elytral intervals 3, 5, 7 and 9 with setae; male Baripus (C.) precordillera was found in the southern area of the protarsomeres without adhesive setae; median Precordillera. This orogenic mountain range is approximately 450 km lobe without ventral mucro, with quadrangular 2008 Ro i g -Ju ñ e n t e t a l .—Tw o Ne w Sp e c i e s o f Ba r i p u s 219

Ta b l e 3. Differences between Baripus (C) mendozensis and Baripus (C.) precordillera.

B. (C.) mendozensis B. (C.) precordillera Adhesive setae of male protarsomeres Small patch Three or four Elytral setae on interval 9 30–38 23–28 Setae of VII sternum 30–41 16–27 Setae mesotibia 6–14 15–20 Male genitalia shape Less curved More curved Apex of median lobe Fig. 5B Fig. 6D Ventral mucro of median lobe Triangular with sharp apex (Figs. 5A, C) Triangular with rounded apex (Figs. 6B–C)

apex; orifice of median lobe with ligula; internal median lobe dorsolateral at least 0.5x length of sac with tooth 6 median lobe 8 3’. Only elytral interval 9 with setae; male protar- 7’. Three setae on each margin of pronotum; male someres with or without small patch of adhesive protarsomeres 1 and 2 with adhesive setae cover- setae; median lobe with ventral mucro; orifice of ing all ventral surface, male protarsomeres sub- median lobe with or without ligula; internal sac quadrangular; sternum VII with fewer than eight without tooth 4 setae; orifice of median lobe lateral 0.33x length of median lobe 12 4 (3). Male protarsomeres 1 and 2 without squamosetae; sternum VII with 30–38 setae; apex of 8 (7). Four setae on each margin of pronotum; median median lobe quadrangular; orifice of median lobe lobe with short and rounded apex; orifice of me- without ligula B. (C.) nevado, new species dian lobe with apical ligula; internal sac with a 4’. Male protarsomeres 1 and 2 with squamosetae complex of fused spiculae, the alpha sclerite; me- on apical region; sternume VII with 16–41 apical dian oviduct joined to bursa copulatrix apically setae; apex of median lobe spatulate; orifice of me- B. (C.) clivinoides (Curtis, 1839) dian lobe with ligula 5 8’. Two setae on each margin of pronotum; median lobe with sharpened apex, orifice of median lobe 5 (4’). Male protarsomeres 1 and 2 with patch of adhe- with or without basal ligula; internal sac without al- sive setae; sternum VII with 30–41 setae; elytral pha sclerite; median oviduct joined to bursa copu- interval 9 with 30–38 setae; first antennomere latrix basally 9 0.8x as long as third, second 0.50x as long as third B. (C.) mendozensis Roig-Juñent 9 (8’). Male protarsomeres without adhesive setae; me- and Cicchino, 1989 dian lobe thin and long, without ventrolateral 5’. Male protarsomeres 1 and 2 with few adhesive grooves; orifice of median lobe long, without setae (three or four); sternum VIII with 16–27 se- ligula; internal sac without tooth tae; elytral interval 9 with 23–28 setae, first anten- B. (C.) flaccus Roig-Juñent and Cicchino, 1989 no-mere as long as third, second 0.33x as long as 9’. Male protarsomeres 1 and 2 with adhesive setae; third. B. (C.) precordillera, new species median lobe with ventrolateral grooves; orifice of median lobe with ligula; internal sac with tooth 6 (3). Lateral margin of pronotum very wide; ligula 10 0.75x length of median lobe orifice; tooth of internal sac slightly sclerotized 10 (9’). Median lobe wide from middle to apex, with deep B. (C.) schajovskoii Roig-Juñent, 1992a grooves; orifice of median lobe with big and wide 6’. Lateral margin of pronotum narrow; ligula 0.50x ligula B. (C.) sulcatipenis Roig-Juñent, 1992a length of median lobe orifice; tooth of internal sac 10’. Median lobe thin throughout its length, grooves sclerotized B. (C.) gentilii Roig-Juñent, 1992a present but weak; orifice of median lobe, with small and thin ligula 11 7 (1’). Two or four setae on each margin of pronotum; male protarsomeres 1 and 2 with or with- 11 (10’). Apex of median lobe thin, extended downwards out small patch of adhesive setae on apical reas a hook gion; sternum VII with 10–24 setae; orifice of B. (C.) deplanatus Roig-Juñent and Cicchino, 1989 220 An n a l s o f Ca r n e g i e Mu s e u m Vo l . 77

11’. Apex rounded, wide The new species B. (C.) nevado and B. (C.) precordil- B. (C.) painensis Roig-Juñent and Cicchino, 1989 lera are included in a well supported clade (76 / 69) by two exclusive characters: 272 (setae of interval 9: 19–29, and 12 (7’). Orifice of median lobe dorsoapical, small, with 411_ (a tooth in the medial region of basal surface of me- ligula; median lobe with shorter apex; internal dian lobe present). The new species is related to the flaccus sac with a complex of unfused sclerotized spicu- clade by two characters: the absence of a tooth in the inter- lae at same position as tooth; female genital tract nal sac (450) and 170 (squamosetae of male protarsomeres without pocket at apex of bursa copulatrix absent). Baripus (C.) nevado and B. (C.) precordillera are B. (C.) neuquensis Roig-Juñent, 1992a related to B. (C.) longitarsis, B. (C.) schajovskoii, B. (C.) 12’. Orifice of median lobe lateroapical, without ligu- gentilii, and B. (C.) mendozensis because of the length of la; median lobe with longer apex; internal sac with antennomeres (82), the disposition of anal setae (302), and tooth; female genital tract with pocket at apex of the presence of ventral fossa in the median lobe (392). bursa copulatrix 13 Within this group, B. (C.) nevado and B. (C.) precordillera conform a monophyletic group with the species of B. 13 (12’). Elytral striae weakly impressed; apex of median (C.) mendozensis, because these are the only three species lobe very long (more than 3x basal width) and within the genus that have a ventral mucro in the median sharpened, apex slender; pocket at apex of bursa 1 copulatrix with narrow base and completely sclero- lobe (41 ). tized; bursa copulatrix ventrally sclerotized 14 The analyses that applied different weights according to the degree of homoplasy yielded the same or a simi- 13’. Elytral striae distinct; apex of median lobe long lar cladogram (Fig. 8) with a fit of 445.6. The cladogram (as least as long as basal width), with rounded differs only in the polytomy of the species B. (C.) minus, apex; pocket at apex of bursa copulatrix wide basally, with a small part sclerotized; bursa copulatrix sclerotized at dorsal region 15 14 (13). Apex of median lobe strongly sharpened; tooth of internal sac very long (about 3x basal width) B. (C.) dentipenis Roig-Juñent, 1992a 14’. Apex of median lobe rounded; tooth of internal sac shorter (length less than three times the width at base) B. (C.) giaii Roig-Juñent, 1992a 15 (13’).Sternum VII with basal setae; tooth of internal sac as long as wide; pocket at apex of bursa copulatrix small (one third spermathecal length) B. (C.) minus Roig-Juñent, 1992a 15’. Sternum VII without basal setae; tooth of internal sac longer than wide; pocket at apex of bursa copulatrix bigger (equal to spermathecal length) B. (C.) chubutensis Roig-Juñent, 1992a

PHYLOGENETIC ANALYSIS Two cladograms of 131 steps were obtained (Fig. 8) from the analyses of the data matrix (Table 2) using equal weights with a consistency index of 0.73 and a retention index of 0.87. In general, support values are high and several clades have values above 80. These results are similar to those obtained previously by Roig-Juñent (1995). However, the phylogenetic relationship of B. (C.) clivinoides is different in this new analysis because it appears as the sister species of the clade (B. (C.) neuquensis, ((B. (C.) minus, B. (C.) chubutensis), (B. (C.) giai, B. (C.) dentipenis))), whereas in the previous analysis it is the adelphotaxon of all the other species of Fig. 7.—Localities inhabited by Baripus precordillera. A, Paramillos; the subgenus Cardiophthalmus. B, Pampa Canota. 2008 Ro i g -Ju ñ e n t e t a l .—Tw o Ne w Sp e c i e s o f Ba r i p u s 221

Fig. 8.—Strict consensus tree, bootstrap / jackknifing support values presented below branches. 222 An n a l s o f Ca r n e g i e Mu s e u m Vo l . 77

the Andean Mountains. One event was the progressive de- sertification of the eastern side of the Andean Mountains, and the other was the formation of high altitude grasslands and other montane habitats in the Andean Mountains. In this study, we focused on some aspects of the distribution patterns of the species of the subgenus Cardiophthal- mus. These species occur in the Patagonian Steppe, one of the dry habitats that resulted from the uplift of the Andean Mountains about 9 mya (Ortiz-Jaureguizar and Cladera 2006). Allopatry and Sympatry of Baripus (Cardiophthalmus).— The species of the subgenus Cardiophthalmus occur mainly in the Patagonian Steppe habitats with the exception of the new species Baripus (C.) nevado that inhabits montane habitats in the Payunia region (Figs. 3A–B), and B. (C.) precordillera that inhabits montane habitats of the Precor- dillera (Figs. 7A–B). This pattern is the same pattern proposed by Noonan (1990, 1992) for the montane carabid beetles of North America. This author proposed a pattern called “northern lowland plus southern mountains” that involves species whose distribution is in lowlands in the north and extends southward along mountains into regions

Fig. 9.—Distribution map: Baripus (Cardiophthalmus) gentilii, Baripus (Cardiophthalmus) mendozensis, Baripus (Cardiophthalmus) nevado, Baripus (Cardiophthalmus) precordillera, and Baripus (Cardiophthalmus) schajovskoii.

B. (C.) chubutensis, and the pair of sister species B. (C.) giaii/ B. (C.) dentipenis.

BIOGEOGRAPHIC CONSIDERATIONS OF BARIPUS Roig-Juñent (1995) proposed a general biogeographic history of the genus Baripus. His phylogenetic analysis suggested that two vicariant events may have separated the three subgenera. This assumption was based on only the current distribution of the genus and the phylogeny at the time, and not on a formal biogeographical analysis (Roig-Juñent 1995). However, the genus was used in other biogeographical analyses together with other taxa from the southern region of South America (Roig-Juñent 1994, Roig-Juñent et al. 2006), and species of this genus were used in analyses of areas of endemism (Morrone et al. 2002, Domínguez et al. 2006). In all of these analyses, three important events that affected the southern biota were identified. The first event was the uplift of the Andean Mountains, which progressively separated the eastern and western slopes and, consequently, isolated the original bio- Fig. 10.—Distribution map: Baripus (Cardiophthalmus) deplanatus, and ta. The other two events were consequences of the uplift of Baripus (Cardiophthalmus) painensis. 2008 Ro i g -Ju ñ e n t e t a l .—Tw o Ne w Sp e c i e s o f Ba r i p u s 223 with warmer or more xeric conditions. For the southern hemisphere, this pattern would be called “Southern Low- land plus Northern Mountains (SLNM)”, given that the southern species of Cardiophthalmus occur in the lowlands, and only two of the northern species occur in mountains. Noonan (1992) described isolation as one of the main factors that promote the evolution of new taxa. He explained that mesic refuges in the mountains during periods of drought may have been important centers for diversification of species. He proposed a model in which the new species of carabids evolved from isolated populations. These populations were previously part of an ancestral species that was widely distributed in the lowlands before the montane-vicariance cycle began. In the SLNM pattern observed for the northern species of Cardiophthalmus, the mountains may have acted as refugia and, later, as centers from which species evolved. Moreover, this group of northern species shows a phylogenetic and biogeographic pattern which is also present in other species of the subgenus that occur exclusively in the southern lowlands. This phylogenetic and biogeographic pattern (PBP) is determined by four characteristics. The first characteristic, which is a phylogenetic relationship, is that the species being compared are sister species. The other three are biogeographical characteristics of the species and are related to their geographic ranges. First, species’ distributions are allopatric. Secondly, one of the species has a larger geographic range than the other. Lastly, the distribution of the more restricted species is peripheral to the distribution of the more widespread species. To assess whether this PBP is real for the species of Cardiophthalmus, we evaluated all sister species within Fig. 11.—Distribution map: Baripus (Cardiophthalmus) chubutensis, the subgenus. The first sister species analyzed includes B. Baripus (Cardiophthalmus) dentipenis, Baripus (Cardiophthalmus) giaii, (C.) mendozensis, B. (C.) precordillera, and B. (C.) nevado. Baripus (Cardiophthalmus) minus, and Baripus (Cardiophthalmus) neuquensis. The distribution pattern of these three sister species shows that B. (C.) mendozensis has a wide distribution, whereas particular species and their sister groups. For example, the B. (C.) nevado and B. (C.) precordillera have restricted dis- species B. (C.) longitarsis, B. (C.) sulcatipenis, and (C.) cliv- tributions and are peripheral to the area of distribution of inoides have allopatric distributions with their sister groups B. (C.) mendozensis (Fig. 4). Another example is the pair (Fig. 8). In addition, B. (C.) minus and B. (C.) chubutensis of sister species B. (C.) schajovskoii and B. (C.) gentilii, have allopatric distributions with the group comprised of which is the sister clade of B. (C.) mendozensis, B. (C.) B. (C.) giaii and B. (C.) dentipenis (Fig. 8), except in one precordillera, and B. (C.) nevado (Fig. 8), and occurs at location where both B. (C.) minus and B. (C.) giaii have more southern latitudes (Fig. 9). In this particular case, B. been recorded (Fig. 11). However, there is a case where (C.) gentilii had a more widespread distribution than B. (C.) the two species B. (C.) neuquensis and B. (C.) minus may schajovskoii, whose distribution is peripheral to the former species. Although these two species have been collected at have a sympatric distribution. We believe that this case of nearby locations, they have never been found in the same sympatry needs to be corroborated. In intensive field stud- site (Fig. 9). Baripus (C.) giaii and B. (C.) dentipenis of ies carried out in western Patagonia, we collected all the which B. (C.) dentipenis is the most widespread, are a pair species mentioned above, and we did not find any two spe- of sister species that have this PBP. However, for this pair of cies of the subgenus Cardiophthalmus coexisting in space species, there is a small area of overlap in their geographic or time. For cases in which these species appear to have ranges because there are two sites where both species were been found in close proximity to one another (mainly data collected (Fig. 11) . Finally, B. (C.) deplanatus and B. (C.) registered in museum collections), it is very possible that painensis form another pair of sister species (Fig. 8) that are the specimens were found in close, but distinct, locations allopatric (Fig. 10) . They do not, however, evince this PBP and were labeled with the same location. because neither species has a restricted distribution. In summary, we found that patterns of distribution Even though they do not have all four characteristics of within Cardiophthalmus are allopatric. Most pairs of sister this PBP, an allopatric distribution was observed between species have a PBP distribution in which one of them has 224 An n a l s o f Ca r n e g i e Mu s e u m Vo l . 77

and B. (C.) nevado have never been collected in the same location. Both species are abundant, conspicuous, and easy to collect. In a transect sampled from the base to the peak of El Nevado Mountain, we found them occurring as close as 7 km from each other but never at the same site. Their distribution pattern may be explained climatic changes during the Cenozoic resulting in changes in the conditions within their area of distribution, leading to the isolation of peripheral populations. Subsequent changes could have led to the current distribution, in which both species occur in close proximity. Something similar may have happened between B. (C.) mendozensis and B. (C.) precordillera, with the difference that their populations are presently far away from each other. When we analyzed the present potential distribution of B. (C.) mendozensis (Fig. 12), we found that this species may be found in areas where it has not yet been recorded, mainly in the southern region of its distribution. Moreover, the analysis showed suitable habitat for B. (C.) mendozensis where the other species described above, B. (C.) nevado, also occurred. This could be because these two species inhabit areas with similar environmental characteristics. However, the potential distribution did not include the areas occupied by the other species. The maximum age

Fig. 12.—Potential distribution of Baripus (Cardiophthalmus) mendozensis. a restricted distribution that is peripheral to the distribution of the more widely distributed species. In the northern- most species, the restricted distribution is associated with mountainous environments. However, this is not the case for the species inhabiting the southern lowlands. Mode of speciation.—The phylogenetic and biogeographic pattern described above led us to infer that the subgenus Cardiophthalmus may have had a peripatric model of speciation. In this model, a small and isolated peripheral population of the widespread species evolves independently of the main population (Mayr 1963; Losos and Glor 2003). The clearest example of this model of speciation is for the two species of the apical clade of B. (C.) mendozensis and B. (C.) nevado. These two species must have been isolated from each other for the speciation process to occur. Considering that the species of Baripus are large-bodied and wingless, they have a low power of dispersal. There- fore, it is possible that fragmentation of the area resulting from physical or climatic barriers may have effectively isolated the two resulting populations. Remarkably, there are today no actual physical nor climatic barriers separating the distributions of these two species, and, in some cases, they Fig. 13.—Potential distribution of Baripus (Cardiophthalmus) occur very close to each other. Baripus (C.) mendozensis mendozensis, Year 2050. 2008 Ro i g -Ju ñ e n t e t a l .—Tw o Ne w Sp e c i e s o f Ba r i p u s 225 of B. (C.) nevado is approximately 5 million years; given an overall decrease in temperatures. However, the model that El Nevado Mountain is that age (Ramos and Folguera, indicated that the current potential distribution is smaller unpublished data). During the 5 million years since the (Fig. 12) than the historic potential distribution, and that end of the Cenozoic, there have been some physiographic the most suitable habitats are currently at higher altitudes. changes in the Payunia region (mainly volcanic activity To further observe the effect of climate change on the in the Quaternary: Bermudez et al. 1993), and significant potential distribution of this species, we used environmen- climatic changes. These changes included fluctuations in tal projections to the year 2050, for which we assumed an temperature and changes in the amount and seasonality of increase of approximately 2° C over the current tempera- precipitation (Ortíz-Jaureguizar and Cladera 2006). Giv- ture. In this scenario, there was a fragmentation of the area en that climatic shifts can cause extinction, splitting, and of distribution of B. (C.) mendozensis, with the most suit- merging of gene pools and clades (Jansson and Dynesius able habitats in the mountains (Fig. 13). These results sup- 2002), we hypothesize that the changes in climatic condi- ported our hypothesis by showing the high sensitivity of B. tions may be the principal cause for changes in the distri- (C.) mendozensis to climatic fluctuations. bution ranges of the ancestors of these species. The changes in the potential distribution range of B. When we modeled the potential distribution of B. (C.) (C.) mendozensis followed the pattern described by Noo- mendozensis using environmental data from 21,000 B.P., nan (1990, 1992) in which mountains are refugia for the we found that the distribution of this species was larger than species inhabiting the lowlands. This phenomenon of re- it is currently (Fig. 14), and that this species was widely traction toward high mountains is the same proposed for distributed in the lowlands. This historic distribution range carabids of the boreal biota, in which the distribution range coincides with an extreme glacial climate episode (dated decreased to include only the highland regions during between 25,000-17,700 B.P. (Moreno, unpublished data)). interglacial periods (warm and dry periods), and the low- Therefore, the ample distribution of B. (C.) mendozensis lands acted as an effective barrier (Ball 1970). in the lowlands could be the result of their avoiding the Therefore, we hypothesize that B. (C.) mendozensis extreme lower temperatures in the highlands because of occupied the larger area in highlands from the north to the south of Mendoza Province on the slopes of the An- dean and extra-Andean mountains. A subsequent climatic change, leading to current conditions, resulted in the isolation of the populations at higher altitudes of the extra-An- dean mountains. These populations speciated resulting in the appearance of B. (C.) precordillera and B. (C.) nevado. Also, the climate change resulted in B. mendozensis inhabiting the lowlands. At present, B. (C.) mendozensis and B. (C.) nevado present close distribution ranges, sharing similar climatic conditions but these species do not coex- ist, perhaps because of interspecific competition. Although there are no similar studies for insects in Argentina, Elgueta (1988) and Covarruvias and Elgueta (1991) proposed a similar model of speciation for species that inhabit the extra-Andean mountain ranges in central Chile. Covarruvias and Elgueta (1991) reject the idea that the Andean Mountains have been a “center of dispersion” from which the species dispersed to the extra-Andean ranges when climatic conditions connected the habitats of these areas, and the insect fauna then diversified by fragmentation and isolation. Elgueta (1988) proposes that the colonizations and disjunctions have been mainly a result of climatic changes from glaciations and that the colonizations occurred from populations of insects that inhabited the lowlands.

CONCLUSIONS The study of regions that have never been explored has allowed us to collect new species of the genus Baripus Fig. 14.—Potential distribution of Baripus (Cardiophthalmus) mendoz- (Cardiophthalmus). These new taxa are the only ones with- ensis, Year 21,000 B.P. in the subgenus that inhabit montane environments, with all 226 An n a l s o f Ca r n e g i e Mu s e u m Vo l . 77

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