''Nepenthes tobaica'' is a tropical

pitcher plant Pitcher plants are several different carnivorous plants which have modified leaves known as pitfall traps—a prey-trapping mechanism featuring a deep cavity filled with digestive liquid. The traps of what are considered to be "true" pitcher p ...

endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...

Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...

. It is particularly abundant around

Lake Toba Lake Toba ( id, Danau Toba) ( Toba Batak: ᯖᯀᯬ ᯖᯬᯅ; romanized: ''Tao Toba'') is a large natural lake in North Sumatra, Indonesia, occupying the caldera of a supervolcano. The lake is located in the middle of the northern part of the ...

, after which it is named. ''Nepenthes tobaica'' is closely related to '' N. angasanensis'', '' N. gracilis'', '' N. mikei'', and '' N. reinwardtiana''.Clarke, C.M. 2001. ''

Nepenthes of Sumatra and Peninsular Malaysia ''Nepenthes of Sumatra and Peninsular Malaysia'' is a monograph by Charles Clarke on the tropical pitcher plants of Sumatra, Peninsular Malaysia, and their minor surrounding islands. It was published in 2001 by Natural History Publications (Bor ...

''. Natural History Publications (Borneo), Kota Kinabalu.

Botanical history

The earliest known collection of ''N. tobaica'' was made by

Johannes Elias Teijsmann Johannes Elias Teijsmann (1 June 1808 – 22 June 1882) was a biologist, botanist and plant collector. He was born in Arnhem, The Netherlands. His surname is sometimes spelled ''Teysmann'', although he himself spelled it ''Teijsmann''. Teijsmann t ...

on February 8, 1856, probably from the

Batak Batak is a collective term used to identify a number of closely related Austronesian ethnic groups predominantly found in North Sumatra, Indonesia, who speak Batak languages. The term is used to include the Karo, Pakpak, Simalungun, Toba, ...

regions. This specimen, which includes female floral material, is H. L. B. 908,155-1106.

Matthew Jebb Matthew Hilary Peter Jebb (born 1958) is an Irish botanist and taxonomist specialising in the ant plant genera ''Squamellaria'', '' Myrmecodia'', '' Hydnophytum'', ''Myrmephytum'' and '' Anthorrhiza'', as well as the carnivorous plant genus '' ...

and

Martin Cheek Martin Roy Cheek (born 1960) is a botanist and taxonomist at the Royal Botanic Gardens, Kew.lectotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes the ...

of ''N. tobaica'' in their 1997 monograph.Jebb, M.H.P. & M.R. Cheek 1997. A skeletal revision of ''Nepenthes'' (Nepenthaceae). ''Blumea'' 42(1): 1–106. ''Nepenthes tobaica'' was formally described in 1928 by

Dutch Dutch commonly refers to: * Something of, from, or related to the Netherlands * Dutch people () * Dutch language () Dutch may also refer to: Places * Dutch, West Virginia, a community in the United States * Pennsylvania Dutch Country People E ...

botanist B. H. Danser in his seminal monograph "

The Nepenthaceae of the Netherlands Indies "The Nepenthaceae of the Netherlands Indies" is a seminal monograph by B. H. Danser on the tropical pitcher plants of the Dutch East Indies and surrounding regions. It was originally published in the ''Bulletin du Jardin Botanique de Buitenzorg'' ...

". At the time, the species was already known from numerous herbarium specimens. Danser suggested a possibly conspecific species in '' N. reinwardtiana'':

"''N. tobaica'' has only been found on the plateau north, east and south of Lake Toba. It is most closely related to ''N. Reinwardtiana'' and I am not quite certain, whether it is perhaps a form of this species, but up till now no intermediate forms have been discovered."

In the scientific literature, ''N. tobaica'' has been confused with the closely related '' N. angasanensis'' on several occasions. ''Nepenthes tobaica'' as described in Danser's 1940 description of '' N. densiflora'' is actually ''N. angasanensis''. Specimens identified as ''N. tobaica'' in Rusjdi Tamin and

Mitsuru Hotta (23 July 1935 – 8 July 2015) was a Japanese botanist best known for his research on Araceae.van Steenis-Kruseman, M.J., ''et al.'' 2006Cyclopaedia of Malesian Collectors: Mitsuru Hotta Nationaal Herbarium Nederland. Hotta was born in Osaka, ...

's 1986 monograph on

n ''Nepenthes'' actually represent both ''N. angasanensis'' and ''N. tobaica''. Some plants sold in the

horticultural Horticulture is the branch of agriculture that deals with the art, science, technology, and business of plant cultivation. It includes the cultivation of fruits, vegetables, nuts, seeds, herbs, sprouts, mushrooms, algae, flowers, seaweeds and no ...

trade under the name ''N. tobaica'' are likely to represent a manmade cross between '' N. khasiana'' and '' N. ventricosa''.

Description

''Nepenthes tobaica'' is a climbing plant. The stem can attain a height of up to 7 m and is up to 6 mm in diameter. Internodes are up to 25 cm long and often round in cross section. However, mature plants may have angular stems because of a groove that originates at the node and extends across most of the internode's length. Neptobaica2

Leaves are

coriaceous This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...

in texture and range from

sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...

to sub- petiolate. The shape of the

lamina Lamina may refer to: Science and technology * Planar lamina, a two-dimensional planar closed surface with mass and density, in mathematics * Laminar flow, (or streamline flow) occurs when a fluid flows in parallel layers, with no disruption betwee ...

, or leaf blade, varies from oblong to

spathulate This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...

. It measures up to 20 cm in length by 4 cm in width. The apical end of the lamina is typically rounded, but may be narrowed and obtuse. Robust plants sometimes have a sub-

peltate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...

tendril insertion. The lamina may be gradually or abruptly contracted towards the amplexicaul base, which clasps the stem for around half of its circumference. One to three longitudinal veins are present on either side of the

midrib This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...

, although they are only prominent in robust specimens.

Pinnate Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and in ...

veins are indistinct.

Tendril In botany, a tendril is a specialized stem, leaf or petiole with a threadlike shape used by climbing plants for support and attachment, as well as cellular invasion by parasitic plants such as ''Cuscuta''. There are many plants that have tendr ...

s grow to 30 cm in length. Rosette and lower pitchers are ovoid in the lower portion and cylindrical above. They are up to 20 cm high by 4 cm wide. A pair of wings runs down the pitcher's ventral surface, often bearing fringe elements either throughout the whole length or only in the upper part. The

gland In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream (endocrine gland) or into cavities inside the body or its outer surface (exocrine gland). Structure De ...

ular region of the pitcher's inner surface is confined to the ovoid portion. A pair of "eye spots" is sometimes present on the inner surface, below the lid attachment. The pitcher mouth is ovate and has an oblique insertion. The narrow

peristome Peristome (from the Greek ''peri'', meaning 'around' or 'about', and ''stoma'', 'mouth') is an anatomical feature that surrounds an opening to an organ or structure. Some plants, fungi, and shelled gastropods have peristomes. In mosses In mosses, ...

(≤5 mm wide) is cylindrical or slightly expanded and bears indistinct teeth. The pitcher lid or operculum is ovate to sub-

orbicular Orbicular is an adjective meaning "circular" Orbicular may also refer to: * Orbicular leaf, a plant leaf shape * Orbicularis oculi muscle, a muscle around the eye * Orbicularis oris muscle, a muscle around the mouth * Orbicular batfish, a specie ...

and has a somewhat

cordate Cordate is an adjective meaning ' heart-shaped' and is most typically used for: * Cordate (leaf shape) The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a ...

base. It lacks appendages. A

spur A spur is a metal tool designed to be worn in pairs on the heels of riding boots for the purpose of directing a horse or other animal to move forward or laterally while riding. It is usually used to refine the riding aids (commands) and to back ...

measuring up to 5 mm in length is inserted near the base of the lid. It may be unbranched,

bifid Bifid refers to something that is split or cleft into two parts. It may refer to: * Bifid, a variation in the P wave, R wave, or T wave in an echocardiogram in which a wave which usually has a single peak instead has two separate peaks * Bifid ci ...

, or trifid. Neptobaica9

Upper pitchers are typically somewhat

infundibular An infundibulum (Latin for ''funnel''; plural, ''infundibula'') is a funnel-shaped cavity or organ. Anatomy * Brain: the pituitary stalk, also known as the ''infundibulum'' and ''infundibular stalk'', is the connection between the hypothalamus and ...

in the lowermost part, becoming narrowly ovoid in the lower third, and finally cylindrical and slightly narrower above. They may be quite large, reaching 25 cm in height by 5 cm in width. The waxy zone of the inner surface is well developed. A pair of remnant ribs is present in place of the ventral wings. The inner portion of the peristome accounts for around 40% of its total cross-sectional surface length.Bauer, U., C.J. Clemente, T. Renner & W. Federle 2012. Form follows function: morphological diversification and alternative trapping strategies in carnivorous ''Nepenthes'' pitcher plants. ''Journal of Evolutionary Biology'' 25(1): 90–102. Other parts of aerial pitchers are similar to their lower counterparts. ''Nepenthes tobaica'' has a

racemose A raceme ( or ) or racemoid is an unbranched, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are produced as the s ...

inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...

. The peduncle and

rachis In biology, a rachis (from the grc, ῥάχις [], "backbone, spine") is a main axis or "shaft". In zoology and microbiology In vertebrates, ''rachis'' can refer to the series of articulated vertebrae, which encase the spinal cord. In this c ...

can each grow to 20 cm in length. Partial peduncles are two-flowered and lack

bract In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...

eoles. In male inflorescences, the

sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coined b ...

s are elliptic-obtuse, being slightly narrower in female ones. Around the town of Prapat, plants have been observed to come into flower in April, although mature

fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...

s are not common at this time.Shivas, R.G. 1984. ''

Carnivorous Plant Newsletter The ''Carnivorous Plant Newsletter'' is the official publication of the International Carnivorous Plant Society (ICPS), the largest such organization in the world. It is headquartered in Walnut Creek, California. History and editorship The newsle ...

'' 13(1): 15–16. ''Nepenthes tobaica'' is characterised by an uneven

indumentum In biology, an indumentum (Latin, literally: "garment") is a covering of trichomes (fine "hairs") on a plant Davis, Peter Hadland and Heywood, Vernon Hilton (1963) ''Principles of angiosperm taxonomy'' Van Nostrandpage, Princeton, New Jersey, pa ...

. Most mature vegetative parts are

glabrous Glabrousness (from the Latin '' glaber'' meaning "bald", "hairless", "shaved", "smooth") is the technical term for a lack of hair, down, setae, trichomes or other such covering. A glabrous surface may be a natural characteristic of all or part of ...

, although the midrib may bear persistent hairs. Groups of white, stellate hairs are often present in the leaf

axil A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...

s. The sepals of this species are densely

tomentose Trichomes (); ) are fine outgrowths or appendages on plants, algae, lichens, and certain protists. They are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae. A covering of any kind of hair on a pl ...

, but the rest of the inflorescence has a sparser covering of short hairs. In some forms, the underside of the lid is a vivid red, making the plants particularly easy to spot amongst other vegetation.

Ecology

''Nepenthes tobaica'' is

. Its natural range was once thought to stretch from the Gayo Lands of

Aceh Aceh ( ), officially the Aceh Province ( ace, Nanggroë Acèh; id, Provinsi Aceh) is the westernmost province of Indonesia. It is located on the northernmost of Sumatra island, with Banda Aceh being its capital and largest city. Granted a s ...

in the north to

Tarutung Tarutung (Dutch: ) is a town and the administrative capital (seat) of North Tapanuli Regency (''Kabupaten Tapanuli Utara''), North Sumatra, Sumatra, Indonesia. in the Batak language means "durian The durian (, ) is the edible fruit of se ...

in the south. Towards the end of the 20th century, new locations of this species were discovered further south. These include Mount Sorik Merapi and a unique high altitude

peat swamp Peat swamp forests are tropical moist forests where waterlogged soil prevents dead leaves and wood from fully decomposing. Over time, this creates a thick layer of acidic peat. Large areas of these forests are being logged at high rates. Peat ...

habitat near Lake Kerinci in

Jambi Jambi is a province of Indonesia. It is located on the east coast of central Sumatra and spans to the Barisan Mountains in the west. Its capital and largest city is Jambi. The province has a land area of 50,160.05 km2, and a sea area of 3, ...

Charles Clarke Charles Rodway Clarke (born 21 September 1950) is a British Labour Party politician, who was the Member of Parliament (MP) for Norwich South from 1997 until 2010, and served as Home Secretary from December 2004 until May 2006. Early life T ...

cites its altitudinal range as 380 to 1800 m

above sea level Height above mean sea level is a measure of the vertical distance (height, elevation or altitude) of a location in reference to a historic mean sea level taken as a vertical datum. In geodesy, it is formalized as ''orthometric heights''. The comb ...

, with a habitat near the

port A port is a maritime facility comprising one or more wharves or loading areas, where ships load and discharge cargo and passengers. Although usually situated on a sea coast or estuary, ports can also be found far inland, such as Ham ...

town of

Sibolga Sibolga (formerly sometimes Siboga) is a city and a port located in the natural harbor of Sibolga Bay on the west coast of North Sumatra province, in Indonesia. It is located on the western side of North Sumatra, facing the Indian Ocean and is ...

constituting the lower extreme and

Mount Pangulubao Mount Pangulubao or Pangulubau ( id, Gunung Pangulubao) is a mountain near Lake Toba in Sumatra. Mount Pangulubao is notable for its large number of native tropical pitcher plant species. These include ''Nepenthes ampullaria'', '' Nepenthes gym ...

the upper extreme. However, according to

and

Martin Cheek Martin Roy Cheek (born 1960) is a botanist and taxonomist at the Royal Botanic Gardens, Kew.

This species typically grows terrestrially in exposed areas. ''Nepenthes tobaica'' commonly grows in forest edges, where it is often sympatric with species of ''

Leptospermum ''Leptospermum'' is a genus of shrubs and small trees in the myrtle family Myrtaceae commonly known as tea trees, although this name is sometimes also used for some species of ''Melaleuca''. Most species are endemic to Australia, with the greate ...

'' and '' Rhodomyrtus''. ''Nepenthes tobaica'' is very abundant in the

region, to the point where it is "difficult to avoid seeing it there". ''N. tobaica'' is also known to grow in high altitude

forest near Lake Kerinci, at an altitude of 1100 m. At this location, the species grows alongside '' N. ampullaria'', '' N. gracilis'', '' N. mirabilis'', '' N. reinwardtiana'', and '' N. spathulata''. The ''Nepenthes'' species and hybrids at this site exhibit high levels of

introgression Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Intr ...

. The vegetation is dominated by species of the genera ''

Rhododendron ''Rhododendron'' (; from Ancient Greek ''rhódon'' "rose" and ''déndron'' "tree") is a very large genus of about 1,024 species of woody plants in the heath family (Ericaceae). They can be either evergreen or deciduous. Most species are nati ...

'' and ''

Melastoma ''Melastoma'' is a genus in the family Melastomataceae. It has over 100 species distributed around Southeast Asia, India, north to Japan, south to Australia and the Pacific Islands. The number of species should probably be reduced according to ...

'', as well as

orchid Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Along with the Asteraceae, they are one of the two largest families of flowering ...

s and

fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...

s. It is very stunted and dense, with few trees exceeding 3 m in height. It is somewhat reminiscent of the highland

heath forest Heath forest is a type of tropical moist forest found in areas with acidic, sandy soils that are extremely nutrient-poor. Notable examples are the Rio Negro campinarana of the Amazon Basin in South America, and the Sundaland heath forests (also kn ...

s around

Bareo Bareo is a small village situated on the National Highway-31 in Nawada District. It shares schools with Namdarganj as other nearby villages in this area. The population consists of both Hindus and Muslims. There is a playground in Bareo which ...

Sarawak Sarawak (; ) is a States and federal territories of Malaysia, state of Malaysia. The largest among the 13 states, with an area almost equal to that of Peninsular Malaysia, Sarawak is located in northwest Borneo Island, and is bordered by the M ...

Borneo Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java, west of Sulawesi, and eas ...

. The

conservation status The conservation status of a group of organisms (for instance, a species) indicates whether the group still exists and how likely the group is to become extinct in the near future. Many factors are taken into account when assessing conservation ...

of ''N. tobaica'' is listed as

Least Concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...

on the

IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...

based on an assessment carried out in 2018.

Pitcher infauna

The pitchers of ''N. tobaica'' play host to a wide variety of infaunal organisms. The

mite Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...

species '' Creutzeria tobaica'' was described from the pitchers of a

Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's List ...

nese ''Nepenthes'' identified as ''N. tobaica''.Fashing, N.J. 2002. ''Australian Journal of Entomology'' 41(1): 7–17. However, ''N. tobaica'' has not been recorded outside of Sumatra.

Related species

In 2001,

performed a

cladistic analysis Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived chara ...

of the ''Nepenthes'' species of Sumatra and

Peninsular Malaysia Peninsular Malaysia ( ms, Semenanjung Malaysia; Jawi: سمننجڠ مليسيا), or the States of Malaya ( ms, Negeri-negeri Tanah Melayu; Jawi: نڬري-نڬري تانه ملايو), also known as West Malaysia or the Malaysian Peninsula, ...

using 70 morphological characteristics of each taxon. The following is a portion of the resultant

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

, showing part of "Clade 6". The sister pair of ''N. angasanensis'' and ''N. mikei'' has 79% support. Neptobaica3

The pitchers of ''N. tobaica'' are similar to those of '' N. adnata'', although the stem and lamina are quite different.

Natural hybrids

Five natural hybrids involving ''N. tobaica'' have been recorded to date; specifically, crosses with '' N. ampullaria'', '' N. reinwardtiana'', '' N. rhombicaulis'', '' N. spathulata'', and '' N. spectabilis''.McPherson, S.R. 2009. ''

Pitcher Plants of the Old World ''Pitcher Plants of the Old World'' is a two-volume monograph by Stewart McPherson on the pitcher plants of the genera ''Nepenthes'' and ''Cephalotus''. It was published in May 2009 by Redfern Natural History Productions and covers all species kno ...

''. 2 volumes. Redfern Natural History Productions, Poole. Dariana 2010
Keanekaragaman ''Nepenthes'' dan pohon inang di Taman Wisata Alam Sicikeh-Cikeh Kabupaten Dairi Sumatera Utara
M.Sc. thesis, University of North Sumatra, Medan.

Notes

:a.The

Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...

description of ''N. tobaica'' from Danser's monograph reads:

''Folia mediocria'' sessilia, lamina lineari-lanceolata, nervis longitudinalibus utrinque 0-1, raro 2, basi lata semiamplexicauli ; ''ascidia rosularum'' ignota ; ''ascidia inferiora'' parte inferiore anguste ovata, medio angustata, os versus infundibuliformia, alis 2 fimbriatis, peristomio operculum versus acuto, cylindrico, 1/2-1 mm lato, costis 1/3-1/4 mm distantibus, saepe indistinctis, dentibus fere 0 ; operculo rotundato-elliptico, facie inferiore plano ; ''ascidia superiora'' parva, e basi infundibuliformi parte inferiore paulum ventricosa, medio leviter angustata, os versus paulum infundibuliformia, costis 2 prominentibus, peristomio operculum versus acuto, cylindrico, 1/2-1 mm lato, costis 1/3-1/4 mm distantibus, saepe indistinctis, dentibus fere 0 ; operculo orbiculari v. rotundato-elliptico facie interiore plana ; ''inflorescentia'' racemus pedicellis 13-15 mm longis fere omnibus 2-floris ; ''indumentum'' in inflorescentiis et ascidiis parcum stellatum, ceterum 0.

:b.In addition to the earliest specimen, collected by Teijsmann in 1856, Danser lists 11 separate collections of ''N. tobaica''.

References

External links

* {{Taxonbar, from=Q3493234 Carnivorous plants of Asia tobaica Endemic flora of Sumatra Plants described in 1928 Taxa named by Benedictus Hubertus Danser