Apistogramma Psammophila – a New Geophagine Dwarf Cichlid (Teleostei: Cichlidae) from the Rio Atabapo Drainage in Colombia and Venezuela

69 (1): 103 –110

Apistogramma psammophila – a new geophagine dwarf cichlid (Teleostei: Cichlidae) from the Rio Atabapo drainage in Colombia and Venezuela

Wolfgang Staeck 1 & Ingo Schindler 2

1 [email protected] — 2 [email protected]

Submitted June 29, 2018. Accepted January 10, 2019. Published online at www.senckenberg.de/vertebrate-zoology on February 15, 2019. Published in print on February 27, 2019. Editor in charge: Uwe Fritz

Abstract Apistogramma psammophila sp. n. is described from the lower Rio Atabapo drainage in Colombia (Departamento Guainía) and Venezuela (Estado Amazonas). It can be distinguished from all congeners by the following combination of characters: Two dark horizontal stripes on ERG\VLGHWZRSHFWRUDOVSRWVVXERUELWDOVWULSHEHFRPLQJPXFKZLGHUYHQWUDOO\DQGQRFDXGDOVSRW,QDGXOWPDOHVVRIWGRUVDODQGDQDO¿QV HDFKZLWKORQJ¿ODPHQWRXVSURORQJDWLRQDOPRVWUHDFKLQJSRVWHULRUPDUJLQRIFDXGDO¿QDQGSHOYLF¿QVZLWK¿ODPHQWRXVH[WHQVLRQQHDUO\ UHDFKLQJPLGGOHRIDQDO¿QEDVH It is most similar to Apistogramma diplotaenia Kullander, 1987, but differs from this species by an irregular pattern of 6 – 8 short vertical dark bar-like markings below the lower lateral stripe, and adult males with reddish area on cheek and chest between eye and pectoral D[LOODDQGGLVWLQFWO\ODQFHRODWHFDXGDO¿Q.

Resumen Apistogramma psammophila sp. n. se describe a partir de la cuenca del río Atabapo bajo en Colombia (Departamento Guainía) y Venezuela (Estado Amazonas). La nueva especie se distingue de todas las demás especies descritas del género Apistogramma por la siguiente com- binación de carácteres diagnosticos: dos bandas laterales oscuras horizontales en el lado del cuerpo, dos manchas pectorales, una banda suborbital que se vuelve ventralmente mucho más ancha, no hay una mancha caudal. En varones adultos aleta dorsal y anal blanda con SURORQJDFLyQ¿ODPHQWRVDODUJDTXHOOHJDKDVWDFDVLHOERUGHSRVWHULRUGHODDOHWDFDXGDO\XQDH[WHQVLyQ¿ODPHQWRVDGHODVDOHWDVSpOYLFDV TXHVHH[WLHQGHKDVWDFDVLODPLWDGGHODEDVHGHODDOHWDDQDO La nueva especie es más similar a Apistogramma diplotaenia Kullander, 1987, pero se difíere de esta especie por un patrón irregular de 6 – OtQHDVRVFXUDVYHUWLFDOHVFRUWDVGHEDMRGHODEDQGDODWHUDOLQIHULRUXQiUHDURML]DHQODPHMLOOD\HOWyUD[HQWUHHORMR\ODD[LODSHFWRUDO en machos adultos y su aleta caudal claramente lanceolada.

Kurzfassung Apistogramma psammophila sp. n. wird aus dem Einzugsbereich des unteren Rio Atabapo in Kolumbien (Departamento Guainía) und Ve- nezuela (Estado Amazonas) beschrieben. Die Art lässt sich von allen anderen beschriebenen Apistogramma-Arten durch die Kombination IROJHQGHUGLDJQRVWLVFKHU0HUNPDOHQXQWHUVFKHLGHQ=ZHLKRUL]RQWDOHGXQNOH6HLWHQElQGHUDXIGHU.|USHUVHLWH]ZHL%UXVWÀHFNHQHLQ VXERUELWDOHU6WUHLIHQGHUYHQWUDOYLHOEUHLWHUZLUGNHLQ6FKZDQ]ZXU]HOÀHFN%HLHUZDFKVHQHQ0lQQFKHQ5FNHQÀRVVHXQG$IWHUÀRVVH EHLQDKHELV]XPKLQWHUHQ(QGHGHU6FKZDQ]ÀRVVHYHUOlQJHUW %DXFKÀRVVHQPLWHLQHUIDGHQI|UPLJHQ(UZHLWHUXQJGLHELVIDVW]XU0LWWH GHU$IWHUÀRVVHQEDVLVUHLFKW Die neue Art ist Apistogramma diplotaenia Kullander, 1987 am ähnlichsten, unterscheidet sich jedoch von dieser durch ein unre- gelmäßiges Muster von 6 – 8 kurzen vertikalen dunklen balkenähnlichen Markierungen unter dem unteren Längsband, einen rötlichen %HUHLFK]ZLVFKHQ$XJHXQGGHP$QVDW]GHU%UXVWÀRVVHQDXI:DQJHXQG%UXVWHUZDFKVHQHU0lQQFKHQXQGGHUHQGHXWOLFKODQ]HWWI|UPLJH 6FKZDQ]ÀRVVH.

Key words Apistogramma diplotaenia, Cichlinae, Departamento Guainía, Estado Amazonas, Geophagini, ichthyology, new species, Orinoco basin, WD[RQRP\.

ISSN 1864-5755 | eISSN 2625-8498 | DOI: 10.26049/VZ69-1-2019-05 103 Staeck, W. & Schindler, I.: Apistogramma psammophila – a new geophagine dwarf cichlid from the Rio Atabapo

Introduction QRO7KHW\SHVDUHGHSRVLWHGLQWKH¿VKFROOHFWLRQVRIWKH Instituto de Ciencias Naturales, Museo de Historia Natu- ral, Universidad Nacional de Colombia in Bogotá (ICN- In 1981 Schmettkamp, a German aquarist, found a single MHN) and in the Senckenberg Naturhistorische Samm- male specimen of an unknown Apistogramma species in OXQJHQ0XVHXPIU7LHUNXQGH'UHVGHQ 07') a shipment of the Cardinal tetra (Paracheirodon axelro- The techniques for taking measurements and meristic di). He called it Doppelband Apistogramma (= Double data follow those described in KULLANDER (1980, 1986). Band Apistogramma) with reference to its unusual color Measurements were made with electronic digital cali- pattern and informally described it as new in the aquari- per to the nearest 0.1 mm. Specimen lengths are given um literature (SCHMETTKAMP, 2QHRIXV :6 FRO- as standard length (SL). Scale rows are numbered as de- lected this species in 1981 and a second time 1986 in the scribed in KULLANDER (1990). Numbers in parentheses Anavilhanas archipelago (lower Rio Negro, Brazil) and DIWHUFRXQWVLQGLFDWHWKHQXPEHURIVSHFLPHQVH[DPLQHG SXEOLVKHGWKH¿UVWGHWDLOVRILWVHFRORJ\ LINKE & STAECK, with that condition. Meristic data are given for eight type 1984, 1994). In 1987, KULLANDER described the species specimens with SL > 20 mm. as Apistogramma diplotaenia from collecting sites in the In the following, we use a pragmatic approach for upper and middle Rio Negro. species recognition (KOTTELAT, 1995). In accordance RÖMER (1992a, b) described reproduction under aqua- ZLWKFXUUHQWWD[RQRPLFSXEOLFDWLRQVRQWKHJHQXVApisto- rium conditions and provided information on the ecologi- gramma (e.g., MESA & LASSO, 2011; VARELLA & SABAJ, FDOFRQGLWLRQVLQWKH¿HOG/DWHUKHGRFXPHQWHGVHYHUDO WKH QHZ VSHFLHV LV GLDJQRVHG E\ H[WHUQDO FKDU- mood-dependent variations in its live coloration, pub- acters. This procedure is consistent with the diagnostic lished further details of its ecology and characterized it variant of the phylogenetic species concept (cf. NIXON & as a sand-dwelling species with a habitat preference of WHEELER, 1990). beaches and sand banks (RÖMER, 1998). Sand-dwelling Comparisons were made with three specimens of contrasts the majority of other members of the genus, Apistogramma diplotaenia (SL 24.5 – 30.5 mm) in the which generally inhabit aggregations of dead leaves. collections of the authors and by consulting published ANDERSON SUREDEO\ZDVWKH¿UVWWRPHQWLRQD sources (KULLANDER, 1987; RÖMER, 1992a, b, 1998). population of A. diplotaenia in the Rio Atabapo (form- ing the border between Columbia and Venezuela). One year later GOTTWALD and KOSLOWSKI (in STAWIKOWSKI et Apistogramma psammophila n. sp. , al. 1995: 62 – 63) published color photos of specimens XUQOVLG]RREDQNRUJDFW$))))'%($ collected in this river. They designated them as Apisto- 49C331082C8C gramma cf. diplotaenia „Orinoco“, based on hybridiza- WLRQH[SHULPHQWVLQGLFDWLQJWKDWSRSXODWLRQVLQWKH5LR )LJV±7DEOH Negro and the Rio Atabapo drainages were probably dif- Holotype. ICN-MHN 24052, male 27.5 mm SL. Colombia (Depar- ferent species. In the following years, a few specimens tamento Guainía), lower Rio Atabapo, close to the mouth of Caño RIWKH$WDEDSRGZDUIFLFKOLGZHUHH[SRUWHGDVDTXDULXP &KDPXFKLQD DSSUR[ƍƍƍ1ƍƍƍ: DOHIWEDQNWULEX- ¿VKIURPERWK&RORPELDDQG9HQH]XHODWR*HUPDQ\ WDU\LPSRUWRIDTXDULXP¿VKWUDGHleg. Javier León Barreto, don. 8QWLOUHFHQWO\H[SRUWVDQGUHSRUWVZHUHUDUHUHQGHU- R. Grossklaus 2018. ing available information about Apistogramma cf. diplo- Paratypes.07')±H[±PP6/9HQH taenia³2ULQRFR´H[WUHPHO\SDWFK\$OWKRXJKWKHGZDUI zuela (Estado Amazonas), lower Rio Atabapo, mouth of an unamed ULJKWEDQN WULEXWDU\ DSSUR[ ƍƍƍ1 ƍƍƍ: VLWXDWHG cichlid from the Rio Atabapo is relatively well known between Isla Zapo and Isla Guaimara (Isla Guanare), import of in the aquarium hobby, it has been neglected by the sci- DTXDULXP¿VKWUDGHleg. and don55LHWVFK07')± HQWL¿FOLWHUDWXUHSXEOLVKHGRQWKH¿VKIDXQDRIWKH2UL- H[±PP6/6DPHGDWDDVLQKRORW\SH noco drainage (KULLANDER, 2003; MESA & LASSO, 2011; ORTEGA-LARA, 7KHRQO\H[FHSWLRQLVLASSO et al. Diagnosis. A slender geophagine dwarf cichlid with (2004). Recent collections and observations at collecting GLVWLQFW VH[XDO GLPRUSKLVP EXW RQO\ VOLJKW VH[XDO GL- sites in the Rio Atabapo revealed previously unknown chromatism. Males grow larger than females and have GHWDLOVRIWKHHFRORJ\RIWKLVVSHFLHVDQGFRQ¿UPHGWKH SURORQJHGUD\VLQWKHLUVRIWGRUVDODQGDQDO¿QDQGD¿OD- hypothesis that it is distinct from the Rio Negro popula- PHQWRXVH[WHQVLRQRIWKH¿UVWSHOYLF¿QUD\ tion. The objective of this paper is to formally describe It can be distinguished from all the other described this dwarf cichlid, distinguish it from A. diplotaenia, and species of Apistogramma by the following combination provide new information on its ecology. of characters: (1) two dark horizontal bands on side of body; (2) two pectoral spots; (3) suborbital stripe becom- ing much wider ventrally; (4) no caudal spot; (5) adult Material and methods PDOHVZLWKVKDOORZGRUVDO¿Q VRIWSDUWRIGRUVDODQG DQDO¿QVHDFKZLWKORQJ¿ODPHQWRXVSURORQJDWLRQDOPRVW UHDFKLQJ SRVWHULRU PDUJLQ RI FDXGDO ¿Q SHOYLF ¿Q 7KHW\SHVSHFLPHQVZHUH¿[HGHLWKHULQHWKDQRORU ZLWK ¿ODPHQWRXV H[WHQVLRQ QHDUO\ UHDFKLQJ PLGGOH RI IRUPDOLQWKHODWWHUVSHFLPHQVWUDQVIHUUHGWRHWKD- DQDO¿QEDVH

104 VERTEBRATE ZOOLOGY — 69 (1) 2019

Fig. 1. Apistogramma psammophila (male, SL 27.5 mm), holotype (ICN-MHN 24052) from Rio Atabapo (Caño Chamuchina, Departa- PHQWR*XDLQtD&ROXPELD RQHGD\DIWHU¿[DWLRQ

Fig. 2. Male of Apistogramma psammophila PP6/ SDUDW\SH 07') IURPORZHU5LR$WDEDSR PRXWKRIDQXQQDPHG ULJKWEDQNWULEXWDU\(VWDGR$PD]RQDV9HQH]XHOD WKUHHZHHNVDIWHU¿[DWLRQ

In general appearance this species is most similar to Table 1. Morphometric data of holotype (ICN-MHN 24052) and Apistogramma diplotaenia Kullander, 1987, but differs ¿YHSDUDW\SHV 07') RIApistogramma psam mo- from this species by 6 – 8 short dark vertical bars of ir- philaLQSHUFHQWRI6/ LQPP PLQ ORZHVWYDOXHPD[ KLJKHVW regular shape in the abdominal region below the lower value, mean = arithmetic mean, sd = standard deviation. lateral band (versus no such pattern), and adult males with reddish area on cheek and chest between eye and pectoral min max mean sd D[LOOD YHUVXVQRVXFKFRORUDWLRQ DQGFDXGDO¿QGLVWLQFWO\ Standard length (mm) 22.8 41.5 32.2 8.25 lanceolate with middle rays longest (versus elongated, Total length 137.5 141.2 139.4 1.36 RYRLGZLWKURXQGHGSRVWHULRUPDUJLQFI)LJ Head length 31.3 32.6 32.1 0.51 Body depth 24.3 27.6 25.5 1.22 Description. 5HIHUWRWKH¿JXUHVIRUJHQHUDODSSHDUDQFH Eye diameter 11.3 13.6 12.3 0.93 Morphometric data are summarized in Table 1. A rela- Interorbital distance 5.7 6.7 6.2 0.45 tively small species of Apistogramma PD[LPXP6/DS- Preorbital length 3.3 3.9 3.6 0.22 SUR[PP %RG\FRPSDUDWLYHO\HORQJDWHG DYHUDJH Peduncle depth 12.4 13.7 13.1 0.47 ERG\GHSWKLQWKHWKUHHODUJHVWVSHFLPHQVRI6/ Peduncle length 14.2 15.0 14.6 0.31 +HDGZLWKDSSUR[LPDWHO\HTXDOO\VWHHSSUHGRUVDODQG 3HFWRUDO¿QOHQJWK 21.8 28.1 25.5 2.61 preventral contours; frontal contour tangential to orbit. 3HOYLF¿QOHQJWK 31.0 49.1 39.8 7.78 Snout short, rounded in lateral and dorsal views. Orbit Length last D spine 16.2 17.1 16.8 0.33 situated in anterior half of head, ventrally reaching into 'RUVDO¿QEDVHOHQJWK 55.6 59.9 58.1 1.90 lower half of head. Mouth terminal, lower jaw slightly $QDO¿QEDVHOHQJWK 18.9 21.8 20.1 1.08 VKRUWHU0D[LOODH[WHQGLQJWRPDUJLQRIRUELW&DXGDOSH- duncle longer than high. 'RUVDO¿QFRPSDUDWLYHO\VKDOORZGRUVDO¿QODSSHWV longation nearly reaching posterior margin of caudal short, with edge rounded or subtruncate in females. In ¿Q$QDO ¿Q ZLWK GLVWDO WLS SRLQWHG H[WHQGLQJ VOLJKWO\ DGXOW PDOHV VRIW GRUVDO ¿Q ZLWK ORQJ ¿ODPHQWRXV SUR- EH\RQGPLGGOHRIFDXGDO¿Q&DXGDO¿QURXQGHGLQIH-

105 Staeck, W. & Schindler, I.: Apistogramma psammophila – a new geophagine dwarf cichlid from the Rio Atabapo

3 4

5 6

7 8

Fig. 3. Adult dominant male of Apistogramma psammophila (no type) from lower Rio Atabapo (right-bank tributary, Estado Amazonas, Venezuela) photographed in aquarium. Fig. 4. Adult female of Apistogramma psammophila (no type) from lower Rio Atabapo (right-bank tributary, Estado Amazonas, Vene- zuela) photographed in aquarium. Fig. 5. Aggressive male of Apistogramma psammophila (no type). Fig. 6. Adult male of Apistogramma psammophila (no type) in neutral mood. Fig. 7. Threatening male of Apistogramma psammophila (no type).

Fig. 8. Adult female of Apistogramma psammophila QRW\SH H[KLELWLQJEURRGFDUHFRORUDWLRQ males, lanceolate in adult males, with middle rays much FHUDWREUDQFKLDOEHDULQJRUXVXDOO\H[WHUQDOJLOOUDN- ORQJHU3HFWRUDO¿QVURXQGHG3HOYLF¿QVZLWK¿ODPHQ- ers. Preopercular serrations variable in number (8 – 17), WRXVSURORQJDWLRQ¿UVWUD\QHDUO\UHDFKLQJPLGGOHRIDQDO irregularly distributed along lower part of vertical limb ¿QEDVH and corner; inconspicuous in smaller specimens (< 20 'RUVDO¿Q;,9 ;9 ;9 $QDO¿Q,,, mm SL). Margin of preopercle smooth in larger speci- E1 row scales 22(6), 23(2). Tube bearing scales 11 – 13 in mens (> 35 mm SL). Lower pharyngeal tooth-plate (dis- upper lateral line and 2 – 6 in lower lateral line. Pectoral sected from one paratype of 41.5 mm SL) wider than ¿QURXQGHGZLWKXVXDOO\UD\V'HQWDOSRUHV)LUVW ORQJ OHQJWK DERXW RI ZLGWK ZLWK WHHWK LQ

106 VERTEBRATE ZOOLOGY — 69 (1) 2019

Fig. 9. Unamed right-hand tributary of the lower Rio Atabapo (Departamento Guainía, Colombia), collecting site of paratypes of Apisto- gramma psammophila.

SRVWHULRUURZDQGWHHWKLQPHGLDQURZ7HHWKELFXV- EHFRPLQJKRUL]RQWDOEHKLQGSHFWRUDOD[LOODDQGFRQYHUJ- pid, two most medial-posterior with prominent major ing with the upper stripe on caudal peduncle and anterior cusp; lateral and rostral teeth slender. HGJHRIFDXGDO¿Q,QWHUVSDFHEHWZHHQWZRVWULSHVJUH\ in males, orange or reddish brown in females. In the Coloration of live specimens )LJV – 8). Description abdominal region below the lower lateral stripe, a pat- based on observations immediately after capture, under- tern of 6 – 8 conspicuous dark vertical bars of different water observations in native habitats and observations of VKDSHVWKH¿UVWEHKLQGWKHSHOYLF¿QRULJLQWKHODVWRQ specimens kept in aquarium. Back, nape and forehead caudal peduncle; interspaces pale and highly contrasted. pale brown. Head pale grey, chest and abdomen whitish. )LQVSDOHJUH\LVKRUEOXLVKPRUHRUOHVVRSDTXH6RIW 7LSVRIGRUVDO¿QODSSHWVUHGIRUPLQJDQDUURZUHGULP portion of dorsal and anal with alternating pattern of RQWKH¿Q,QDGXOWPDOHVUHGGLVKDUHDRQFKHHNDQGFKHVW brownish dots and hyaline windows on posterior mem- EHWZHHQH\HDQGSHFWRUDOD[LOOD EUDQHV 'RUVDO OREH RI FDXGDO ¿Q ZLWK SDWWHUQ RI GDUN Manifestation and intensity of black or dark brown dots alternating with hyaline windows; ventral half hya- markings is variable and mood-dependent as in other line. species of Apistogramma (see RÖMER, 1998; VARELLA & SABAJ, 2014). Suborbital stripe running obliquely from Geographical distribution and ecology. Apistogramma postero-ventral margin of orbit across cheek; stripe not psammophila is currently known only from localities in uniform, antero-dorsal portion narrower than pupil, pos- the lower Rio Atabapo, and is possibly endemic to this tero-ventral portion much wider, often enlarged as con- drainage. One of the documented collecting sites is situ- spicuous black blotch. Postorbital stripe intense, con- DWHG FORVH WR WKH PRXWK RI &DxR &KDPXFKLQD DSSUR[ WLQXRXVZLWKGDUNVWULSHRQÀDQNV3UHRUELWDOVWULSHOHVV ƍƍƍ1ƍƍƍ: DOHIWEDQNWULEXWDU\$QRWKHU GLVWLQFW3HFWRUDOD[LOODZLWKWZREODFNLVKSHFWRUDOVSRWV is in the mouth of an unnamed right-bank tributary (ap- one dorsally, the other ventrally on adjacent side; also a SUR[ ƍƍƍ1 ƍƍƍ: VLWXDWHG EHWZHHQ ,VOD EODFNLVKVSRWDERYHXSSHUEDVHRISHFWRUDO¿Q7ZRGDUN Zapo and Isla Guaimara (Isla Guanare). horizontal stripes on side of body side. The upper one All specimens of the type material were collected in DERYHORZHUODWHUDOOLQHVWUDLJKWZLWKDSSUR[LPDWHO\ the low-water season between the second half of Janu- half the depth of a scale; lower stripe slightly wider, DU\DQGWKH¿UVWKDOIRI)HEUXDU\'XULQJWKLVWLPHWKH width nearly the depth of one scale; Lower stripe anteri- ZDWHU OHYHO ZDV DERXW VL[ PHWHUV EHORZ LWV PD[LPXP orly curved, running ventrad from the postorbital stripe, height in the high-water season, but still more than one

107 Staeck, W. & Schindler, I.: Apistogramma psammophila – a new geophagine dwarf cichlid from the Rio Atabapo

10 11

Fig. 10. Male Apistogramma diplotaenia from the Anavilhanas archipelago (lower Rio Negro, Brazil).

Fig. 11. Threatening male of Apistogramma diplotaenia (docu- PHQWDWLRQRIURXQGHGFDXGDO¿Q

12 Fig. 12. )HPDOH RI Apistogramma diplotaenia H[KLELWLQJ EURRG care coloration. meter above its minimum. Both collecting sites are sandy Etymology. 7KHVSHFLHVHSLWKHW *UHHNȥȐȝȝȠȢ VDQG EHDFKHV )LJ 7KH ¿VK ZHUH H[FOXVLYHO\ IRXQG DQG ĳȚȜȓĮ IULHQGVKLS LVDQDGMHFWLYH FISVDPPRSKLORXV FROOHFWHGDWDGLVWDQFHRIVHYHUDOPHWHUVIURPWKHH[SRVHG It refers to the habitat preference of the species. EHDFKZKHUHWKHZDWHUGHSWKYDULHGIURPDSSUR[LPDWHO\ 1.5 to at least 2 meters. According to our underwater ob- VHUYDWLRQVLQWKH¿HOGWKHGZDUIFLFKOLGVOLYHGVLQJO\DQG Discussion spread far apart over a wide sandy area where shelter was scarce. Both collecting sites are typical blackwater habitats Apistogramma diplotaenia and A. psammophila have a ZLWK FOHDU YHU\ DFLGLF DQG H[WUHPHO\ VRIW WHDFRORUHG vicariant distribution in the Rio Negro and Rio Atabapo, water. The following data were collected at both locali- respectively. Several other pairs of closely related cich- ties in different years: pH 4.1 – PD[LPXPHOHFWULFDO lid species have a similar complementary distribution FRQGXFWLYLW\ 6FP WRWDO DQG WHPSRUDU\ KDUGQHVV pattern in those two drainages (KULLANDER & FERREIRA, OG+ GHJUHHVRIKDUGQHVV ZDWHUWHPSHUDWXUH – 1988; STAWIKOWSKI, 1989). &HOVLXV Allopatric sister species may pose problems because morphology and osteological differences are often small Reproductive biology. According to our aquarium ob- DQG KHQFH GLVFULPLQDWLRQ DQG FODVVL¿FDWLRQ LV GLI¿FXOW servations, spawning takes place in concealed localities. (MAYR, 1969). Closely related species in South Ameri- $SSUR[LPDWHO\WRHJJVDUHDWWDFKHGWRWKHFHLOLQJ can cichlids are usually most easily distinguished by dif- of the shelter. The female guards the eggs, larvae, and ferences in live color patterns (KULLANDER, 1980, 1989; free-swimming young while the male defends the terri- KULLANDER & FERREIRA, 1988). WRU\:KHQNHSWLQDQDTXDULXPWKH¿VKIRUPSDLUVDW Even minor deviations in coloration or patterns of least during the breeding season. GDUNPDUNLQJVPD\EHRIELRORJLFDOVLJQL¿FDQFHDVWKHVH During brood care, females in many species in Apisto- differences are important in mate choice. Cichlid spe- grammaVSHFLHVH[KLELWVSHFLDOFRORUSDWWHUQVFRQVLVWLQJ cies are separated mostly by pre-zygotic and pre-mating, RIVSHFLHVVSHFL¿FLQWHQVHEODFNPDUNLQJVRQDFRQWUDVW- EHKDYLRUDO LVRODWLQJ PHFKDQLVPV PRUH VSHFL¿FDOO\ E\ ing bright yellow background. However, A. psammoph- female choice preferences (KOCHER, 2004). In cichlids, ilaODFNVVXFKVSHFLDOL]HGEURRGFDUHFRORUDWLRQ )LJ mating preferences and mate choice are based on visual Only the color of the anterior interspace between the two factors (COULDRIDGE & ALEXANDER, 2002; BLAIS et al., ODWHUDOVWULSHVLVLQWHQVL¿HGDQGWXUQVRUDQJHRUUHGGLVK 2009), and the females of Apistogramma species can dis- brown during brood care. FULPLQDWHEHWZHHQFRQVSHFL¿FDQGKHWHURVSHFL¿FFRXUW-

108 VERTEBRATE ZOOLOGY — 69 (1) 2019 ing males even if they are closely related and look similar Acknowledgements (BEISENHERZ et al., 2006; READY et al., 2006; ENGELKING et al., 2010). There are indications that such pre-mating behavioral mechanisms isolate Apistogramma diplotae- :H DUH LQGHEWHG WR -DYLHU /HyQ %DUUHWR -RVp /HRQHO &DOGHUyQ nia and A. psammophila (see GOTTWALD and KOSLOWSKI Alonso Rodrigo Sotomayor (Puerto Inírida, Colombia), the staff of )DXQD$TXiWLFD$PD]RQDV 3XHUWR$\DFXFKR9HQH]XHOD DQGWKH STAWIKOWSKI , in et al. 1995: 62 – 63). friendly people of the Communidade Santa Rosa (Departamento Among congeneric species, Apistogramma psam- Guainía, Colombia) for valuable information or administrative mophila resembles only A. diplotaenia2QWKHLUÀDQNV DVVLVWDQFH GXULQJ ¿HOGZRUN LQ &RORPELD DQG 9HQH]XHOD:H DUH the two species have the same unique color pattern of two obliged to Roland Rietsch (Berlin, Germany) and René Grossklaus dark horizontal bands that fuse both anteriorly in front (Merenschwand, Switzerland) for providing the type material. :HWKDQN8ZH)ULW] 07' DQGWKH,QVWLWXWRGH&LHQFLDV1DWX- RIWKHSHFWRUDO¿QDQGSRVWHULRUO\RQWKHFDXGDOSHGXQ- rales, Museo de Historia Natural (ICN-MHN) in Bogotá for depos- FOH)HPDOHVODFNWKH\HOORZEUHHGLQJFRORUDWLRQW\SLFDO LWLQJWKHW\SHPDWHULDOLQWKHLULQVWLWXWHV:HDUHJUDWHIXOWR0DUN of other ApistogrammaVSHFLHV FI)LJVDQG ,Q Sabaj for his valuable suggestions to improve an earlier version of addition, both are sand-dwelling species, an ecological WKHWH[WDQGDOVRWKDQNWKUHHDQRQ\PRXVUHYLHZHUVIRUWKHLUFULWLFDO specialization that is unique within the genus. comments. KULLANDER (1987) regarded A. diplotaenia as a highly distinctive species in the genus. Despite the thorough comparative osteological study, placement of A. diplo- References taenia within the genus remained obscure, and there- fore, KULLANDER did not hypothesize close relatives. A ANDERSON, T. B. (1994). Ecuador und Kolumbien 1993. DCG-In- preliminary phylogenetic analysis based on molecular formationen, 25, 203 – 216. data (MILLER & SCHLIEWEN, 2005) produced similar re- BEISENHERZ, W., RÖMER, U., MÜLLER, M. & ENGELKING, B. (2006). VXOWV7KLVVWXG\FRQ¿UPHGA. diplotaenia as a distinct :LHZlKOHQApistogramma :HLEFKHQLKUH3DUWQHU",QWUDXQG LQWHUVSH]L¿VFKH3DUWQHUZDKOGXUFK:HLEFKHQYRQApistogram- lineage, but its relationship to congeners remained un- ma cacatuoides Hoedeman, 1951 (Teleostei, Perciformes, Cich- solved. On the basis of the results of a cluster analysis lidae), pp. 7 – 8 in: GREVEN, H. & RIEHL, R. (eds): Biologie der based on phenotypical and behavioral characters, RÖMER $TXDULHQ¿VFKH. Velten, Tetra Verlag. (2006) placed Apistogramma diplotaeniaLQDFRPSOH[ BLAIS, J., PLENDERLEITH, M., RICO, C., TAYLOR, M. I., SEEHAUSEN, O., OOSETERHOUT, C. TURNER, G. F. of its own within his Apistogramma agassizii supercom- VAN & (2009). Assortative mat- LQJDPRQJ/DNH0DODZLFLFKOLG¿VKSRSXODWLRQVLVQRWVLPSO\ SOH[ predictable from male nuptial colour. BMC Evolutionary Bio- Apistogramma species live under different environ- logy, 9, 53. mental conditions in a variety of water bodies such as COULDRIDGE, V. C. K. & ALEXANDER, G. J. (2002). Color patterns rivers, lakes, lagoons, brooks, creeks, ponds, and even and species recognition in four closely related species of Lake seasonal pools, which may dry up during the low-water Malawi cichlids. Behavior Ecology, 13, 59 – 64. ENGELKING, B., RÖMER, U. & BEISENHERZ, W. ,QWUDVSHFL¿F season. In spite of this, their habitat preferences are sur- colour preference in mate choice by female Apistogramma ca- prisingly uniform and homogeneous. They prefer lentic catuoides Hoedeman, 1951 (Teleostei, Perciformes, Cichlidae). ZDWHUVDQGOLYHFORVHWRWKHPDUJLQVLQH[WUHPHO\VKDOORZ Vertebrate Zoology, 60, 123 – 138. zones where the water depth generally varies from ap- KOCHER, T. D. $GDSWLYHHYROXWLRQDQGH[SORVLYHVSHFLDWLRQ SUR[LPDWHO\WHQWRIRUW\FHQWLPHWHUV7KHW\SLFDOELRWRSH 7KHFLFKOLG¿VKPRGHONature Reviews Genetics, 5, 288 – 298. KOTTELAT, M. (1995). Systematic studies and biodiversity: The need of the vast majority of Apistogramma species is the leaf for a pragmatic approach. Journal of Natural History, 29, 565 – litter found at the edge of small forest streams or lagoons. 569. Only a few species prefer shelters in aquatic plants or KULLANDER62 $WD[RQRPLFDOVWXG\RIWKHJHQXVApisto- ÀRDWLQJPHDGRZVRIPDFURSK\WHV HJEichhornia spp., gramma Regan, with a revision of Brazilian and Peruvian spe- Cabomba spp. or Paspalum repens) near the water’s sur- cies (Teleostei: Percoidei: Cichlidae). Bonner Zoologische Mo- nographien, 14, 1 – 152. face (STAECK & SCHINDLER, 2008). KULLANDER, S. O. (1986). &LFKOLG¿VKHVRIWKH$PD]RQ5LYHUGUDLQ- 8QOLNH WKHLU FRQVSHFL¿FV Apistogramma psam- age of Peru. Stockholm, Swedish Museum of Natural History. mophila and A. diplotaenia live in completely different KULLANDER, S. O. (1987). A new Apistogramma species (Teleostei, habitats. They occupy an ecological niche that is unique Cichlidae) from the Rio Negro in Brazil and Venezuela. Zoo- within the genus (cf. RÖMER, IRUZHGLGQRW¿QGA. logica Scripta, 16, 259 – 270. KULLANDER, S. O. (1989). Description of a new Acaronia species psammophila in shallow waters or leaf litter close to the (Teleostei, Cichlidae) from the Rio Orinoco and Rio Negro ULYHUEDQN,QFRQWUDVWWRLWVFRQVSHFL¿FVA. psammophila drainages. Zoologica Scripta, 18, 447 – 452. lives over sandy bottoms at a distance of several meters KULLANDER, S. O. (1990). Mazarunia mazarunii (Teleostei: Cichli- IURPWKHH[SRVHGEHDFKZKHUHWKHZDWHULVPRUHWKDQ dae), a new genus and species from Guyana, South America. RQHPHWHUGHHS )LJ Ichthyological Exploration of Freshwaters, 1, 3 – 14. KULLANDER, S. O. (2003). Cichlidae, pp. 605 – 654 in: REIS, R. E., 6\VWHPDWLFV DQG LQWHUVSHFL¿F SK\ORJHQHWLF UHODWLRQ- KULLANDER, S. O. & FERRARIS, C. J. (eds) Check List of the ships in Apistogramma, the second largest genus of neo- Freshwater Fishes of South and Central America. Porto Ale- tropical cichlids, need further analysis. gre, Edipucrs.

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KULLANDER, S. O. & FERREIRA, E. J. G. (1988). A new Satanoperca READY, J. S., SAMPAIO, I., SCHNEIDER, H., VINSON, C., DOS SANTOS, species (Teleostei, Cichlidae) from the Amazon River basin in T. & TURNER, G. F. (2006). Colour forms of Amazonian cichlid Brazil. Cybium, 12, 343 – 355. ¿VKUHSUHVHQWUHSURGXFWLYHO\LVRODWHGVSHFLHV Journal of Evo- LASSO, C. A., MOJICA, J. I., USMA, J. S., MALDONADO-OCAMPO, J. A., lutionary Biology, 19, 1139 – 1148. DONASCIMIENTO, C., TAPHORN, D. C., PROVENZANO)LASSO- RÖMER, U. (1992a). Beobachtungen zur Aquarienbiologie von Api- ALCALÁ, O. M., GALVIS, G., VÁSQUEZ, L., LUGO, M., MACHA- sto gramma diplotaenia Kullander, 1987. DCG-Informationen, DO-ALLISON, A., ROYERO, R., SUÁREZ, C. & ORTEGA-LARA, A. 23, 166 – 170. (2004). Peces de la cuenca del río Orinoco. Parte 1: Lista de RÖMER, U. E )UHLODQGEHREDFKWXQJHQ DQApistogramma di- especies y distribución por subcuencas. Biota Colombiana, 5, plo taenia Kullander, 1987. Buntbarsch-Jahrbuch, 1, 58 – 71. 95 – 158. RÖMER, U. (1998). Cichlid Atlas, volume 1. Melle, Mergus Verlag. LINKE, H. & STAECK, W. (1984). Amerikanische Cichliden I. Kleine RÖMER, U. (2006). Cichlid Atlas, volume 2. Melle, Mergus Verlag. Buntbarsche. Melle, Tetra Verlag. SCHMETTKAMP, W. (1981). Neues aus der Apistogramma-Szene: 4. LINKE, H. & STAECK, W. (1994). American Cichlids I. Dwarf Cich- Apistogramma spec. nov. – Doppelband-Apistogramma. DCG- lids. Melle, Tetra-Press. Informationen, 12, 148 – 149. MAYR, E. (1969). Principles of Systematic Zoology. New York, STAECK: SCHINDLER, I. (2008). Apistogramma erythrura sp. n. – McGraw-Hill. a new geophagine dwarf cichlid (Teleostei: Cichlidae) from MESA, S. L. M. & LASSO, C. A. (2011). Revisión del género Apisto- the río Mamoré drainage in Bolivia. Vertebrate Zoology, 58, gramma Regan, 1913 (Perciformes, Cichlidae) en la cuenca 197 – 206. del río Orinoco. Serie Editorial Recursos Hidrobiológicos y STAWIKOWSKI, R. (1989). Ein neuer Cichlide aus dem oberen Orino- Pesqueros Continentales de Colombia III. Bogotá, Instituto co-Einzug: Uaru fernandezyepezi n. sp. (Pisces: Perciformes: Humboldt. Cich lidae). Bonner zoologische Beiträge, 40, 19 – 26. MILLER, M. & SCHLIEWEN, U. (2005). The molecular phylogeny of STAWIKOWSKI, R., KOSLOWSKI, I. & BOHNET, V. (eds, 2005). Süd ame- the genus Apistogramma – a working hypothesis, pp. 22 – 25 in: rikanische Zwergcichliden/South American Dwarf Cichlids. STAWIKOWSKI, R., KOSLOWSKI, I. & BOHNET, V. (eds) Südameri- Stuttgart (DATZ Sonerheft), Eugen Ulmer Verlag, 129 pp. kanische Zwergcichliden/South American Dwarf Cichlids. Stutt - VARELLA, H. R. & SABAJ PÉREZ, M. H. (2014). A titan among dwarfs – gart, DATZ Sonderheft, Eugen Ulmer. Apistogramma kullanderi, new species (Teleostei: Cichlidae). NIXON, K. C. & WHEELER,4' $QDPSOL¿FDWLRQRIWKHSK\ Ichthyological Exploration Freshwaters, 25, 243 – 258. logenetic species concept. Cladistics, 6, 211 – 223. ORTEGA-LARA, A. (2016). Guía visual de los principales peces ornamentales continentales de Colombia. Santiago de Cali, AUNAP.

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