跟著城市嚮導「老臺北胃」，用味道認識臺北

很多朋友來臺北，
都會問我同一個問題：
「臺北小吃那麼多，到底該從哪裡開始吃？」
夜市裡攤位一字排開、老店藏在巷弄轉角，
看起來都很有名，卻又怕吃錯、踩雷，
結果行程走完，反而沒真正記住臺北的味道。
我常被朋友笑說是「老臺北胃」。
不是因為特別會吃，而是因為在這座城市待久了，
知道哪些味道是陪著臺北人成長的日常。
這篇文章，就是我整理的一份清單。
如果你第一次來臺北，
我會帶你從這 10 樣最具代表性的臺北小吃開始，
不追一時爆紅、不走浮誇路線，
而是讓你吃完後能真正理解
原來，這就是臺灣的小吃文化。
跟著老臺北胃走，
用最簡單的方式，
把臺北的味道，一樣一樣記在心裡。

我怎麼選出這 10 大臺北小吃？

在臺北，
你隨便走進一條夜市或老街，
都可以輕易列出 30 種以上的小吃。
所以這份清單，
不是「臺北最好吃」的排名，
 而是我站在「第一次來臺北的旅客」角度，
做的推薦。
身為一個被朋友稱作「老臺北胃」的人，
我選這 10 樣小吃時，心裡一直放著幾個原則。

一吃就知道：這就是臺灣味

燒烤、火鍋很好吃，
但換個城市、換個國家，也吃得到。
我挑的，是那種
只要一入口，就會讓人聯想到的臺灣味。
 不需要解釋太多，舌頭就能懂。

不只是好吃，而是有「臺北日常感」

臺北的小吃迷人，
不只在味道，
而在它融入生活的方式。
我在意的是：

1. 會不會出現在早餐、宵夜、下班後
2. 有沒有陪伴這座城市很久的記憶

吃完之後，你會記得臺北

最後一個標準很簡單。
如果你回到家，
還會突然想起某個味道、某碗熱湯、某個攤位的香氣
那它就值得被放進這份清單裡。

接下來的 10 樣臺北小吃，
就是我會親自帶朋友去吃的在地美食。
不趕行程、不拚數量，
而是一口一口，
慢慢認識臺北。

第 1 家：饌堂－黑金滷肉飯（雙連店）｜一碗就懂臺灣人的日常

如果只能用一道料理，
 來解釋臺灣人的日常飲食，
 那我一定會先帶你吃滷肉飯。
在臺北，滷肉飯不是什麼特別的節慶料理，
 而是從早餐、午餐到宵夜，
 默默陪著很多人長大的味道。
而在眾多滷肉飯之中，
饌堂－黑金滷肉飯（雙連店），
 我很常帶第一次來臺北的朋友造訪的一家。

為什麼第一站，我會選饌堂？
饌堂的滷肉飯，走的是**「黑金系」路線**。
滷汁顏色深、香氣厚，
卻不死鹹、不油膩。
滷肉切得細緻，
肥肉入口即化，搭配熱騰騰的白飯，
每一口都是很完整、很臺灣的味道。
對第一次吃滷肉飯的旅客來說，
這種風味夠經典、也夠穩定，
不需要太多心理準備，就能理解為什麼臺灣人這麼愛它。

不只是好吃，而是「現在的臺北感」
饌堂並不是那種躲在深巷裡的老攤，
空間乾淨、節奏俐落，
卻沒有失去滷肉飯該有的靈魂。
這也是我會推薦給旅客的原因之一：
它保留了臺灣小吃的核心味道，
同時也讓第一次來臺北的人，
吃得安心、坐得舒服。

老臺北胃的帶路小提醒
如果是第一次來：

1. 一定要點招牌黑金滷肉飯
   
2. 可以加一顆滷蛋，風味會更完整
3. 搭配簡單的小菜，就很有臺灣家常感

這不是那種吃完會驚呼「哇！」的料理，
而是會讓你在幾口之後，
慢慢理解
原來，臺灣人的日常，就是這樣被一碗飯照顧著。

地址：103臺北市大同區雙連街55號1樓

電話：0225501379

菜單：https://bio.site/ZhuanTang

第 2 家：富宏牛肉麵｜臺北深夜也醒著的一碗熱湯

如果說滷肉飯代表的是臺灣人的日常，
 那牛肉麵，
 就是很多臺北人心中最有份量的一餐。
而在臺北提到牛肉麵，
 富宏牛肉麵，
 幾乎是夜貓族、加班族、外地旅客一定會被帶去的一站。

為什麼老臺北胃會帶你來吃富宏？
富宏最讓人印象深刻的，
不是華麗裝潢，
而是那鍋永遠冒著熱氣的紅燒湯頭。
湯色濃而不混，
帶著牛骨與醬香慢慢熬出的厚度，
喝起來溫潤、不刺激，
卻會在嘴裡留下很深的記憶點。
牛肉給得大方，
燉到軟嫩卻不鬆散，
搭配彈性十足的麵條，
每一口都很直接、很臺北。

不分時間，任何時候都適合的一碗麵
富宏牛肉麵最迷人的地方，
在於它陪伴了無數個臺北的夜晚。
不管是深夜下班、看完演唱會、
或是剛抵達臺北、還沒適應時差，
這裡總有一碗熱湯在等你。
對旅客來說，
這種不用算時間、不用擔心打烊的安心感，
本身就是一種臺北特色。

老臺北胃的帶路小提醒
第一次來富宏，我會這樣點：

1. 紅燒牛肉麵是首選
   
2. 如果想吃得更過癮，可以加點牛筋或牛肚
3. 湯先喝一口原味，再視情況調整辣度

這不是精緻料理，
卻是一碗能在任何時刻撐住你的牛肉麵。
在臺北，
很多夜晚，
就是靠這樣一碗熱湯走過來的。

地址：108臺北市萬華區洛陽街67號

電話：0223713028

菜單：https://www.facebook.com/pages/富宏牛肉麵-原建宏牛肉麵/

第 3 家：士林夜市・吉彖皮蛋涼麵｜臺北夏天最有記憶點的一口清爽

如果你在夏天來到臺北，
 一定會很快發現一件事
 這座城市，真的很熱。
也正因為這樣，
 臺北的小吃世界裡，
 才會出現像「涼麵」這樣的存在。
而在士林夜市，
 吉彖皮蛋涼麵，
 就是我很常帶旅客來吃的一家。

為什麼在夜市，我會帶你吃涼麵？
很多人對夜市的印象，
都是炸物、熱湯、重口味。
但真正的臺北夜市，
其實也很懂得照顧人的胃。
吉彖的涼麵，
冰涼的麵條拌上濃郁芝麻醬，
再加上切得細緻的皮蛋，
入口的第一瞬間，
就是一種「被降溫」的感覺。
那種清爽，
不是沒味道，
而是在濃香與清涼之間取得剛剛好的平衡。

皮蛋，是靈魂，也是臺灣味的關鍵
對很多外國旅客來說，
皮蛋是既好奇、又有點猶豫的存在。
但我常說，
如果要嘗試皮蛋，
涼麵是一個非常溫柔的起點。
芝麻醬的香氣會先接住味蕾，
皮蛋的風味則在後段慢慢出現，
不衝、不嗆，
反而多了一層深度。
很多人吃完後，
都會露出那種「原來是這樣啊」的表情。

老臺北胃的帶路小提醒
第一次點吉彖皮蛋涼麵，我會建議：

1. 一定要選皮蛋款，才吃得到特色
2. 醬料先拌勻，再吃，風味會更完整
3. 如果天氣真的很熱，這一碗會救你一整晚

這不是華麗的小吃，
卻非常臺北。
在悶熱的夜晚，
站在夜市人潮裡，
吃著一碗涼麵，
你會突然明白——

原來臺北的小吃，連氣候都一起考慮進去了。

地址：111臺北市士林區基河路114號

電話：0981014155

菜單：https://www.facebook.com/profile.php?id=100064238763064

第 4 家：胖老闆誠意肉粥｜臺北人深夜最踏實的一碗粥

如果你問我，
 臺北人在深夜、下班後，
 最容易感到被安慰的食物是什麼——
 我會毫不猶豫地說：肉粥。
而提到肉粥，
 胖老闆誠意肉粥，
 就是很多老臺北人口中的那一味。

為什麼這一碗粥，會被叫做「誠意」？
胖老闆的肉粥，看起來很簡單。
白粥、肉燥、配菜，
沒有華麗擺盤，也沒有複雜作法。
但真正坐下來吃，你會發現：
這碗粥，不敷衍任何一個細節。
粥體滑順、不稀薄，
肉燥香而不膩，
搭配各式家常小菜，
一口一口吃下去，
很自然就會放慢速度。
這種味道，
不是要你驚艷，
而是要你安心。

這不是觀光小吃，而是臺北人的生活片段
胖老闆誠意肉粥，
最迷人的地方，
就是它的客人。
你會看到：

1. 剛下班的上班族
2. 熬夜後來吃一碗熱粥的人
3. 熟門熟路、點菜不用看菜單的老客人

這些畫面，
比任何裝潢都更能說明這家店在臺北的位置。
對旅客來說，
這是一個走進臺北人日常的入口。

老臺北胃的帶路小提醒
第一次來吃，我會這樣建議：

1. 肉粥一定要點，這是主角
2. 配幾樣小菜一起吃，才有完整體驗
3. 不用急，慢慢吃，這碗粥就是要你放鬆

這不是為了拍照而存在的小吃，
而是那種
**會讓人記得「那天晚上，我在臺北吃了一碗很溫暖的粥」**的味道。

地址：10491臺北市中山區長春路89-3號

電話：0913806139

菜單：https://lin.ee/xxbYZyS

第 5 家：圓環邊蚵仔煎｜夜市裡最不能缺席的臺灣經典

如果要選一道
 最常出現在旅客記憶裡的臺灣小吃，
 蚵仔煎一定排得上前幾名。
而在臺北，
 圓環邊蚵仔煎，
 就是那種很多臺北人從小吃到大的存在。

為什麼蚵仔煎，這麼能代表臺灣？
蚵仔煎的魅力，
不在於精緻，
而在於它把幾種看似簡單的食材，
煎成了一種獨特的口感。
新鮮蚵仔的海味、
雞蛋的香氣、
地瓜粉形成的滑嫩外皮，
最後再淋上甜中帶鹹的醬汁，
一口下去，
就是夜市的完整畫面。
這種味道，
很難在其他國家找到替代品。

圓環邊，吃的是記憶感
圓環邊蚵仔煎，
沒有多餘的包裝，
也不刻意迎合潮流。
它留下來的原因很簡單
味道夠穩、節奏夠快、
讓人一吃就知道「對，就是這個」。
對旅客來說，
這是一家
不需要研究、不需要比較，就能安心點蚵仔煎的地方。

老臺北胃的帶路小提醒
第一次吃蚵仔煎，我會這樣建議：

1. 趁熱吃，口感最好
   
2. 不用急著加辣，先吃原味
3. 醬汁是靈魂，別急著把它拌掉

蚵仔煎不是細嚼慢嚥的料理，
它屬於人聲鼎沸、鍋鏟作響的夜市時刻。
站在人群裡，
吃著一盤熱騰騰的蚵仔煎，
你會很清楚地感受到
這，就是臺北的夜晚。

地址：103臺北市大同區寧夏路46號

電話：0225580198

菜單：https://oystera.com.tw/menu

第 6 家：阿淑清蒸肉圓｜第一次吃肉圓，就該從這裡開始

說到臺灣小吃，
 很多人腦中一定會出現「肉圓」兩個字。
但真正吃過之後才會發現，
 肉圓，從來不只有一種樣子。
在臺北，
 阿淑清蒸肉圓，
 就是我很常拿來介紹「清蒸派肉圓」的一家。

清蒸肉圓，和你想像的不一樣
不少旅客對肉圓的第一印象，
來自油炸版本，
外皮厚、口感重。
而阿淑的清蒸肉圓，
完全是另一個方向。
外皮晶瑩、滑嫩，
帶著自然的彈性，
不油、不膩，
一入口反而顯得清爽。
內餡扎實，
豬肉香氣清楚，
搭配特製醬汁，
味道層次簡單卻很乾淨。

為什麼我會推薦給第一次來臺北的旅客？
因為這顆肉圓，
不需要適應期。
它不刺激、不厚重，
即使是第一次嘗試臺灣小吃的人，
也能輕鬆接受。
對旅客來說，
這是一顆
「吃得懂、也記得住」的肉圓。

老臺北胃的帶路小提醒
第一次來阿淑，我會這樣吃：

1. 直接點一顆清蒸肉圓，吃原味
2. 醬汁先別全部拌開，邊吃邊調整
3. 放慢速度，感受外皮的口感變化

這不是夜市裡熱鬧喧囂的料理，
而是那種
安靜地展現臺灣小吃功夫的味道。
當你吃完這顆肉圓，
會更明白一件事
臺灣小吃的魅力，
往往藏在這些細節裡。

地址：242新北市新莊區復興路一段141號

電話：0229975505

第 7 家：胡記米粉湯｜一碗最貼近臺北早晨的味道

如果說前面幾樣小吃，
 是臺北的熱鬧與記憶，
 那麼米粉湯，
 就是這座城市最真實的日常。
而在臺北，
 胡記米粉湯，
 是很多人從小吃到大的存在。

為什麼米粉湯，這麼「臺北」？
米粉湯不是重口味料理，
它靠的不是刺激，
而是一碗清澈卻有深度的湯。
胡記的湯頭，
用豬骨慢慢熬出香氣，
喝起來清爽、不油，
卻能在喉嚨留下溫度。
米粉細軟，
吸附湯汁後入口順滑，
簡單到不能再簡單，
卻正是臺北人習以為常的早晨風景。

配菜，才是這一碗的靈魂延伸
在胡記吃米粉湯，
主角雖然是湯，
但真正讓人滿足的，
往往是那些小菜。
紅燒肉、豬內臟、燙青菜，
隨意點上幾樣，
湯一口、菜一口，
就是很多臺北人記憶中的早餐組合。
對旅客來說，
這是一種
不需要解釋，就能融入的臺北生活感。

老臺北胃的帶路小提醒
第一次來胡記，我會這樣建議：

1. 一定要點米粉湯，湯先喝
2. 再配 1～2 樣小菜，體驗會完整很多
3. 這一餐適合慢慢吃，不用趕

這不是為了觀光而存在的小吃，
而是一碗
每天準時出現在臺北人生活裡的湯。
當你坐在店裡，
聽著湯勺碰撞的聲音，
你會突然感覺到——
原來，臺北的早晨，
就是從這樣一碗米粉湯開始的。

地址：106臺北市大安區大安路一段9號1樓

電話：0227212120

第 8 家：藍家割包｜一口咬下的臺灣街頭記憶

如果要選一道
 外國旅客一看到就會好奇、吃完又會記住的小吃，
 割包，一定在名單裡。
而在臺北，
 藍家割包，
 就是我很放心帶旅客來認識這道經典的一站。

割包，為什麼被叫做「臺灣漢堡」？
割包的結構其實很簡單：
鬆軟的白饅頭、
燉得入味的滷五花肉、
酸菜、花生粉、香菜。
但真正迷人的，
是這些元素組合在一起時，
形成的層次感。
肉香、甜味、鹹味、清爽度，
在一口之間同時出現，
沒有誰搶戲，
卻彼此剛好。
這種平衡感，
正是臺灣小吃很迷人的地方。

藍家割包不是走浮誇路線，
它給人的感覺很直接
就是你期待中的割包樣子。
饅頭柔軟不乾，
五花肉肥瘦比例恰到好處，
入口即化卻不膩口，
花生粉的甜香收尾，
讓整體味道非常完整。
對第一次吃割包的旅客來說，
這是一個
不會出錯、也很容易愛上的版本。

老臺北胃的帶路小提醒
第一次吃藍家割包，我會這樣建議：

1. 直接點招牌割包，不要改配料
2. 如果有香菜，建議保留，味道會更完整
3. 趁熱吃，饅頭口感最好

割包不是精緻料理，
卻非常有記憶點。
站在街頭，
拿著一顆熱騰騰的割包，
邊走邊吃，
你會很清楚地感受到
這一口，就是臺灣的街頭生活。

地址：100臺北市中正區羅斯福路三段316巷8弄3號

電話：0223682060

菜單：https://instagram.com/lan_jia_gua_bao?utm_medium=copy_link

第 9 家：御品元冰火湯圓｜臺北夜晚最溫柔的一碗甜

吃了一整天的臺北小吃，
 到了這個時候，
 胃其實已經差不多滿了。
但只要天氣一涼，
 或夜色慢慢降下來，
 你還是會想找一碗——
 不是為了吃飽，而是為了舒服的甜點。
這時候，我通常會帶你來 御品元冰火湯圓。

為什麼叫「冰火」？這碗湯圓的關鍵就在這裡
御品元最有特色的地方，
就在於它的「冰火交錯」。
熱騰騰的湯圓，
外皮軟糯、內餡濃香，
搭配冰涼清甜的桂花蜜湯，
一口下去，
溫度在嘴裡交替出現。
不是衝突，
而是一種很細膩的平衡。
這樣的吃法，
也正是臺灣甜點很擅長的地方——
不張揚，但很有記憶點。

這是一碗，會讓人慢下來的甜點
和夜市裡熱鬧的甜品不同，
御品元的冰火湯圓，
更像是一個讓人停下腳步的存在。
你會發現，
坐在這裡吃湯圓的人，
說話聲都會不自覺地變小。
對旅客來說，
這不只是吃甜點，
而是一個
把白天的熱鬧慢慢收進回憶裡的時刻。

老臺北胃的帶路小提醒
第一次吃御品元，我會這樣建議：

1. 點招牌冰火湯圓，體驗完整特色
2. 先單吃湯圓，再搭配湯一起吃
3. 放慢速度，這一碗不適合趕時間

這不是為了拍照而存在的甜點，
而是一碗
會讓你記得「那天晚上在臺北，很舒服」的湯圓。

地址：106臺北市大安區通化街39巷50弄31號

電話：0955861816

菜單：https://instagram.com/lan_jia_gua_bao

第 10 家：頃刻間綠豆沙牛奶專賣店｜把臺北的味道，留在最後一口清甜

走到這一站，
 其實已經不需要再吃什麼大份量的東西了。
這時候，
 最適合的，
 是一杯不吵鬧、不張揚，
 卻會默默留在記憶裡的飲品。
頃刻間綠豆沙牛奶，
 就是我很常用來替一天畫下句點的選擇。

綠豆沙牛奶，為什麼這麼「臺灣」？
在臺灣，
飲料不只是解渴，
而是一種生活節奏。
綠豆沙牛奶看起來簡單，
但真正好喝的版本，
靠的是火候、比例，
還有耐心。
頃刻間的綠豆沙，
口感細緻、不粗顆，
甜度自然、不膩口，
牛奶的加入，
讓整杯變得柔順而溫和。
這不是衝擊味蕾的飲料，
而是一種
喝完之後，會覺得剛剛那一刻很舒服的甜。

為什麼我會用它當作最後一站？
因為它很臺北。
你可以外帶，
邊走邊喝；
也可以站在店門口，
慢慢把杯子喝空。
沒有儀式感，
卻很真實。
對旅客來說，
這杯綠豆沙牛奶，
就像是把今天吃過的所有味道，
溫柔地整理好，
帶走。

老臺北胃的帶路小提醒
第一次喝頃刻間，我會這樣建議：

1. 直接點招牌綠豆沙牛奶
   
2. 正常甜就很剛好，不用特別調整
3. 找個角落慢慢喝，別急著趕路

這一杯，
不會讓你驚呼，
卻會在回程的路上，
突然想起來。
原來，臺北的味道，是這樣結束一天的。

地址：111臺北市士林區小北街1號

電話：0228818619

菜單：https://instagram.com/chill_out_moment?igshid=YmMyMTA2M2Y=

如果只有 3 天的自助旅行在臺北，怎麼吃這 10 家？

第一次來臺北，
時間有限、胃容量也有限，
與其每一家都趕，不如照著節奏吃。
這份 3 天小吃路線，
是老臺北胃會帶朋友實際走的版本：
不爆走、不硬塞，
讓你每天都吃得剛剛好。

臺北 3 天小吃推薦行程表（老臺北胃版本）

天數	時段	店家名稱	小吃類型
Day 1	午餐	饌堂－黑金滷肉飯（雙連店）	滷肉飯
Day 1	下午	阿淑清蒸肉圓	肉圓
Day 1	晚餐	富宏牛肉麵	牛肉麵
Day 1	宵夜	胖老闆誠意肉粥	粥品
Day 2	早餐	胡記米粉湯	米粉湯
Day 2	下午	藍家割包	割包
Day 2	晚上	士林夜市－吉彖皮蛋涼麵	涼麵
Day 2	夜市	圓環邊蚵仔煎	蚵仔煎
Day 3	下午	御品元冰火湯圓	甜點
Day 3	收尾	頃刻間綠豆沙牛奶專賣店	飲品

雖然每個小吃的地點都有一點距離，但是你也知道，好吃的小吃，是值得你花一點時間前往品嘗
老臺北胃的小提醒

1. 不需要每一家都點到最滿
2. 留一點餘裕，才會想再回來
3. 臺北小吃的魅力，不在於吃多少，而在於記住了什麼味道
   

當你照著這 3 天走完，
你會發現，
臺北不是靠一兩道名菜被記住的，
而是靠這些看似日常、卻很真實的小吃。
下次再來，老臺北胃再帶你吃更深的那一輪。

老臺北胃帶路｜這 10 口，就是我心中的臺北

寫到這裡，
 其實已經不是在推薦哪一家小吃了。
而是在回頭看，
 這座城市，是怎麼用食物陪著人生活的。
滷肉飯、牛肉麵、肉粥、米粉湯，
 不是為了成為觀光名單而存在，
 而是每天默默出現在臺北人的日子裡。
夜市裡的蚵仔煎、涼麵、割包，
 熱鬧、吵雜、節奏很快，
 卻也正是臺北最真實的樣子。
而最後那碗湯圓、那杯綠豆沙牛奶，
 則是在一天結束時，
 替味蕾留下一個溫柔的句點。

如果你問我，
「這 10 家是不是臺北最好吃的小吃？」
我會說，
它們不一定是排行榜第一名，
卻是我真的會帶朋友去吃的版本。
因為它們吃得到：

1. 臺北人的日常
2. 巷弄裡的熟悉感
3. 不需要解釋，就能被理解的味道

如果你是第一次來臺北，
跟著這份清單走，
你不一定會吃得最飽，
但你一定會記得——
臺北，是什麼味道。
而如果有一天，
你又再回到這座城市，
走進熟悉的街口、
看到冒著熱氣的小攤，
你也會開始懂得，
為什麼老臺北胃，
總是記得這些看似平凡的滋味。
因為，真正留在心裡的，
從來不是吃過多少，
而是哪一口，讓你想起臺北。

胖老闆誠意肉粥價格合理嗎？

走完這 10 家，

你可能會發現一件事胡記米粉湯吃起來順口嗎？

臺北的小吃，其實不急著被你記住。

它們就安靜地存在在街角、夜市、轉彎處，胖老闆誠意肉粥推薦必點嗎？

等你有一天，再回到這座城市。富宏牛肉麵在地人怎麼說？

如果你是第一次來臺北，胖老闆誠意肉粥份量有誠意嗎？

希望這份「老臺北胃帶路」的清單，

能幫你少一點猶豫、多一點安心。

不用擔心踩雷，藍家割包真的推薦嗎？

也不用為了排行而奔波，頃刻間綠豆沙牛奶專賣店份量有誠意嗎？

只要照著節奏走，

你就會吃到屬於自己的臺北味道。

而如果你已經來過臺北，

那更希望這篇文章，士林夜市－吉彖皮蛋涼麵當正餐適合嗎？

能帶你走進那些

你可能錯過、卻一直都在的日常小吃。

因為真正迷人的旅行，

從來不是把清單全部打勾，

而是某一天，

你突然想起那碗飯、那口湯、那杯甜，藍家割包好吃嗎？

然後在心裡對自己說一句：阿淑清蒸肉圓冬天適合吃嗎？

「下次再去臺北，還想再吃一次。」

把這篇文章存起來、分享給一起旅行的人，

或是在規劃行程時，再回來看看。

讓味道，成為你認識臺北的方式。

下一次來臺北，

別急著走遠。

老臺北胃，藍家割包好吃嗎？

會一直在這些地方，

等你再回來。

Yash Sondhi went looking for differences in color vision between two closely related moths, one of which is active at night and the other during the day. Instead, he found differences in the way they kept time. Credit: Jeremy Squire Researchers discovered a unique form of speciation in moths, driven by differences in activity times and supported by genetic changes in their clock genes. This study highlights the evolution of moths through temporal isolation, providing a concrete example of speciation at the molecular level. How does one species become two? If you’re a biologist, that’s a loaded question. The consensus is that, in most cases, the process of speciation occurs when individuals from a single population become geographically isolated. If they remain separate long enough, they lose the ability to interbreed. A new study published in the journal Proceedings of the Royal Society B: Biological Sciences demonstrates what happens when a less common form of speciation occurs. Rather than being separated by a physical barrier, such as a mountain range or an ocean, members of a species can become separated in time. The Case Study of Moths The researchers focused on two closely related moth species with overlapping ranges in the southeastern United States. “These two are very similar,” said lead author Yash Sondhi, who conducted research for the study while working at Florida International University and later at the Florida Museum of Natural History. “They’ve differentiated along this one axis, which is when they fly.” Rosy maple moths, in the genus Dryocampa, look like what you’d get if Roald Dahl painted something from a fever dream. They bear a thick lion’s mane above their head and abdomen, and their vibrant scales are the color of strawberry and banana taffy. Both male and female rosy moths fly exclusively at night. As their name implies, rosy maple moths often use maple trees as host plants. Their caterpillars can be so numerous and voracious that they sometimes strip away the leaves on a tree. Credit: Jeremy Squire Pink-striped oakworm moths, in the genus Anisota, are less flashy, with subtle grades of ochre, umber, and marl. While females of this species are active at dusk and early evening, the males prefer to fly during the day. Sondhi knew from previous research that these two groups, Dryocampa and Anisota, originated from a single species approximately 3.8 million years ago, which is relatively recent on evolutionary time scales. There’s a handful of species in the genus Anisota, all of which are active during the day. The nocturnal rosy maple moths are the only species in the genus Dryocampa. Genetic Insights from Moth Species Sondhi specializes in the biology of insect vision and saw the moth pair as the perfect opportunity to explore how vision evolves when a species switches up its pattern of activity. But things didn’t go as planned. “I went in looking for differences in color vision. Instead, we found differences in their clock genes, which in hindsight makes sense,” Sondhi said. Clock genes control the circadian rhythm of plants and animals. The ebb and flow of the proteins they create causes cells to become either active or dormant over a period of roughly 24 hours. They affect everything from metabolism and cell growth to blood pressure and body temperature. For any organism reversing its pattern of activity, clock genes are virtually guaranteed to be involved. “It’s a system that’s been retained in everything from fruit flies to mammals and plants. They all have some kind of time-keeping mechanism,” he said. Investigating Genetic Expressions Sondhi compared the transcriptomes of the two moths. Unlike genomes, which contain the entirety of an organism’s DNA, transcriptomes contain only the subset of genetic material that is being actively used to make proteins. This makes them useful for exploring differences in protein levels throughout the day. As expected, Sondhi found a number of genes that were expressed in different quantities in the two moth species. Nocturnal rosy maple moths invested more energy in their sense of smell, whereas the day-flying oakworm moth produced more genes associated with vision. Discoveries in Gene Expression There were, however, no differences in the genes that confer the ability to see color. That doesn’t necessarily mean that their color vision is identical, but if differences do exist, they are likely at the level of tuning and sensitivity and not in the structure of the genes themselves. There was an additional gene that stood out. Disconnected, or disco, was expressed at different levels during the day and night in both species. In fruit flies, disco is known to indirectly influence circadian rhythms through the production of neurons that transmit clock enzymes from the brain to the body. The disco gene Sondhi found in his moth samples was twice the size of its fruit fly counterpart, and it had additional zinc fingers — active portions of a gene that directly interacts with DNA, RNA and proteins. It seemed likely that changes in the disco gene were at least partially responsible for the switch to night-flying in rosy maple moths. When he compared the disco gene of rosy maple moths with the one in oakworms, he found 23 mutations that made each distinct from the other. The mutations were also located in active portions of the gene, meaning they likely contribute to observable physical differences between the moths. Sondhi was looking at evolution in action. “If this is functionally confirmed, this is a really concrete example of the mechanism behind how they speciated at the molecular level, which is rare to come by,” he said. Implications for Evolutionary Biology The study is also an important push for a better understanding of the various ways in which life sustains and propagates itself. When genetics first became a field of study, researchers focused most their efforts on a few representative species, such as fruit flies or lab mice. This was done primarily for the sake of expediency, but it limits how much we know about broad biological patterns. Just as a human is not a lab mouse, a moth is not a fruit fly. “As species continue to decline due to climate change and other anthropogenic changes, we’ll need to genetically engineer a greater number of the ones that remain to enable drought tolerance, for example, or to be active in light polluted regimes. To do that consistently, having a broader pool of functionally characterized genes across organisms is crucial. We can’t just use Drosophila,” Sondhi said. Reference: “Day–night gene expression reveals circadian gene disco as a candidate for diel-niche evolution in moths” by Yash Sondhi, Rebeccah L. Messcher, Anthony J. Bellantuono, Caroline G. Storer, Scott D. Cinel, R. Keating Godfrey, Andrew J. Mongue, Yi-Ming Weng, Deborah Glass, Ryan A. St Laurent, Chris A. Hamilton, Chandra Earl, Colin J. Brislawn, Ian J. Kitching, Seth M. Bybee, Jamie C. Theobald and Akito Y. Kawahara, 1 August 2024, Proceedings B. DOI: 10.1098/rspb.2024.0591

This schematic shows the difference in length between a PCFG gray whale born in 2020 vs one born prior to the year 2000. OSU researchers determined that a full-grown PCFG gray whale born in 2020 is expected to reach an adult body length that is 1.65 meters (about 5 feet, 5 inches) shorter than a gray whale born prior to 2000. For PCFG gray whales that grow to be 38-41 feet long at full maturity, that accounts for a loss of more than 13% of their total length. Credit: Courtesy K.C. Bierlich, OSU Marine Mammal Institute A study from Oregon State University shows that gray whales off the Pacific Northwest have shrunk significantly since 2000, indicating potential broader environmental issues and underscoring the urgent need for further research into the factors affecting their decline. A recent study by Oregon State University has revealed that gray whales, which feed in the shallow coastal waters of the Pacific Northwest during the summer, have experienced a notable reduction in body length since approximately the year 2000. The smaller size could have major consequences for the health and reproductive success of the affected whales, and also raises alarm bells about the state of the food web in which they coexist, researchers say. “This could be an early warning sign that the abundance of this population is starting to decline, or is not healthy,” said K.C. Bierlich, co-author on the study and an assistant professor at OSU’s Marine Mammal Institute in Newport. “And whales are considered ecosystem sentinels, so if the whale population isn’t doing well, that might say a lot about the environment itself.” The study, published in Global Change Biology, looked at the Pacific Coast Feeding Group (PCFG), a small subset of about 200 gray whales within the larger Eastern North Pacific (ENP) population of around 14,500. This subgroup stays closer to shore along the Oregon coast, feeding in shallower, warmer waters than the Arctic seas where the bulk of the gray whale population spends most of the year. Recent studies from OSU have shown that whales in this subgroup are smaller and in overall worse body condition than their ENP counterparts. The current study reveals that they’ve been getting smaller in recent decades. Technological Advances in Whale Measurement The Marine Mammal Institute’s Geospatial Ecology of Marine Megafauna (GEMM) Lab has been studying this subgroup of gray whales since 2016, including flying drones over the whales to measure their size. Using images from 2016-2022 of 130 individual whales with known or estimated age, researchers determined that a full-grown gray whale born in 2020 is expected to reach an adult body length that is 1.65 meters (about 5 feet, 5 inches) shorter than a gray whale born prior to 2000. For PCFG gray whales that grow to be 38-41 feet long at full maturity, that accounts for a loss of more than 13% of their total length. If the same trend were to happen in humans, that would be like the height of the average American woman shrinking from 5 feet, 4 inches to 4 feet, 8 inches tall over the course of 20 years. “In general, size is critical for animals,” said Enrico Pirotta, lead author on the study and a researcher at the University of St. Andrews in Scotland. “It affects their behavior, their physiology, their life history, and it has cascading effects for the animals and for the community they’re a part of.” Drone image of two gray whales off the Oregon coast. Credit: Courtesy OSU Marine Mammal Institute Whale calves that are smaller at weaning age may be unable to cope with the uncertainty that comes with being newly independent, which can affect survival rates, Pirotta said. For adult gray whales, one of the biggest concerns is reproductive success. “With them being smaller, there are questions of how effectively these PCFG gray whales can store and allocate energy toward growing and maintaining their health. Importantly, are they able to put enough energy toward reproduction and keep the population growing?” Bierlich said. Scarring on PCFG whales from boat strikes and fishing gear entanglement also makes the team concerned that smaller body sizes with lower energy reserves may make the whales less resilient to injuries. Environmental Impacts on Food Sources The study also examined the patterns of the ocean environment that likely regulate food availability for these gray whales off the Pacific coast by tracking cycles of “upwelling” and “relaxation” in the ocean. Upwelling sweeps nutrients from deeper to shallower regions, while relaxation periods then allow those nutrients to remain in shallower areas where light allows for the growth of plankton and other tiny organisms, including the prey of gray whales. “Without a balance between upwelling and relaxation, the ecosystem may not be able to produce enough prey to support the large size of these gray whales,” said co-author Leigh Torres, associate professor and director of the GEMM Lab at OSU. The data show that whale size declined concurrently with changes in the balance between upwelling and relaxation, Pirotta said. “We haven’t looked specifically at how climate change is affecting these patterns, but in general we know that climate change is affecting the oceanography of the Northeast Pacific through changes in wind patterns and water temperature,” he said. “And these factors and others affect the dynamics of upwelling and relaxation in the area.” Now that they know the PCFG gray whales’ body size is declining, researchers say they have a lot of new questions about the downstream consequences of that decline and the factors that could be contributing to it. “We’re heading into our ninth field season studying this PCFG subgroup,” Bierlich said. “This is a powerful dataset that allows us to detect changes in body condition each year, so now we’re examining the environmental drivers of those changes.” Reference: “Modeling individual growth reveals decreasing gray whale body length and correlations with ocean climate indices at multiple scales” by Enrico Pirotta, K. C. Bierlich, Leslie New, Lisa Hildebrand, Clara N. Bird, Alejandro Fernandez Ajó and Leigh G. Torres, 07 June 2024, Global Change Biology. DOI: 10.1111/gcb.17366 The other co-authors on the paper were Lisa Hildebrand, Clara Bird, and Alejandro Ajó at OSU and Leslie New at Ursinus College in Pennsylvania.

Attenborough’s long-beaked echidna, photographed by a camera trap. Credit: Expedition Cyclops An international team rediscovered the rare Attenborough’s long-beaked echidna in Indonesia, also uncovering new species and a cave system. These findings, achieved with local community collaboration, contribute significantly to biodiversity and geological research. A long-beaked echidna named after Sir David Attenborough and last seen by scientists in 1961 has been photographed for the first time in an Indonesian tropical forest. An international team of researchers worked with local communities to deploy over 80 camera traps to film the elusive animal. Besides rediscovering the echidna, the team uncovered a wealth of species completely new to science, including beetles, spiders, and a remarkable tree-dwelling shrimp. Rediscovery of Attenborough’s Long-Beaked Echidna More than sixty years after it was last recorded, an expedition team has rediscovered an iconic, egg-laying mammal in one of the most unexplored regions of the world. Attenborough’s long-beaked echidna, named after famed broadcaster Sir David Attenborough, was captured for the first time in photos and video footage using remote trail cameras set up in the Cyclops Mountains of Indonesia’s Papua Province. Diverse Discoveries Amidst Challenges Alongside the echidna’s rediscovery, the expedition – a partnership between the University of Oxford, Indonesian NGO Yayasan Pelayanan Papua Nenda (YAPPENDA), Cenderawasih University (UNCEN), Papua BBKSDA, and the National Research and Innovation Agency of Indonesia (BRIN), Re:Wild – made many other remarkable finds. These included Mayr’s honeyeater, a bird lost to science since 2008; an entirely new genus of tree-dwelling shrimp; countless new species of insects; and a previously unknown cave system. This was despite the difficulties posed by extremely inhospitable terrain, including venomous animals, blood-sucking leeches, malaria, earthquakes, and exhausting heat. Attenborough’s long-beaked echidna, filmed using a camera trap. Credit: Expedition Cyclops One of the World’s Most Unusual Mammals Finally Caught On Film Recorded by science only once in 1961, Attenborough’s long-beaked echidna is a monotreme: an evolutionarily distinct group of egg-laying mammals that includes the platypus. This echidna species is so special because it is one of only five remaining species of monotremes, the sole guardians of this remarkable branch of the tree of life. Echidnas are notoriously difficult to find since they are nocturnal, live in burrows, and tend to be very shy. Attenborough’s long-beaked echidna has never been recorded anywhere outside the Cyclops Mountains, and is currently classified as Critically Endangered on the IUCN Red List of Threatened Species A temporary expedition campsite on the northern slopes of the Cyclops Mountains. Members of the Yongsu Sapari community helped prepare paths and campsites for the expedition to conduct their research. Credit: Expedition Cyclops To give themselves the best chance of finding one, the team deployed over 80 trail cameras, making multiple ascents of the mountains, and climbing more than 11,000 meters (more than the height of Everest) in the process. For almost the entire four weeks that the team spent in the forest, the cameras recorded no sign of the echidna. On the last day, with the last images on the final memory card, the team obtained their shots of the elusive mammal – the first-ever photographs of Attenborough’s echidna. The identification of the species was later confirmed by Professor Kristofer Helgen, mammalogist and chief scientist and director of the Australian Museum Research Institute (AMRI). The Echidna’s Distinctive Features Dr. James Kempton, a biologist from the University of Oxford who conceived of and led the expedition, said: “Attenborough’s long-beaked echidna has the spines of a hedgehog, the snout of an anteater, and the feet of a mole. Because of its hybrid appearance, it shares its name with a creature of Greek mythology that is half human, half serpent. The reason it appears so unlike other mammals is because it is a member of the monotremes – an egg-laying group that separated from the rest of the mammal tree-of-life about 200 million years ago.” Expedition leader Dr. James Kempton (University of Oxford) collecting specimens after ascending the slopes of the Cyclops Mountains. Credit: Expedition Cyclops “The discovery is the result of a lot of hard work and over three and a half years of planning,” he added. “A key reason why we succeeded is because, with the help of YAPPENDA, we have spent years building a relationship with the community of Yongsu Sapari, a village on the north coast of the Cyclops Mountains. The trust between us was the bedrock of our success because they shared with us the knowledge to navigate these treacherous mountains, and even allowed us to research on lands that have never before felt the tread of human feet.” About Attenborough’s long-beaked echidna: Attenborough’s long-beaked echidna, Zaglossus attenboroughi, is not known to live outside the Cyclops Mountains and biologists still have many unanswered questions about its habitat and ecology. Attenborough’s long-beaked echidna is an EDGE species: a threatened species that has few close relatives on the evolutionary tree of life. They have evolved independently of other mammals for about 200 million years. The echidna has cultural significance for the people of Yongsu Sapari, who have lived on the northern slopes of the Cyclops Mountains for eighteen generations. When there is a conflict within the community, rather than fighting, there is a tradition that one party goes up into the Cyclops to search for an echidna while the other party goes to the ocean to find a marlin. Both creatures were so difficult to find that it would often take decades or a whole generation to locate them, but, once found, the animals symbolized the end of the conflict and a return to harmonious relationships in the village. The echidna has only been scientifically recorded once, when it was discovered by Pieter van Royen – a Dutch botanist – in 1961. Since then it has only been known from reports of sightings by the Yongsu Sapari community, and indirect signs during pre-expedition work in 2022. These signs, also observed during the expedition, included “nose pokes,” holes in the ground left by echidnas after using their long, slightly curved snouts to probe for underground invertebrates. A Treasure Trove of Discoveries Alongside searching for the echidna, the expedition carried out the first comprehensive assessment of invertebrate, reptile, amphibian, and mammal life in the Cyclops Mountains. With the support of local guides in the expedition team, the scientists were able to create makeshift labs in the heart of the jungle with benches and desks made from forest branches and vines. A new species of terrestial shrimp, found in the soil and in the trees of the Cyclops Mountains. This shrimp belongs to a lineage that is normally found on the seashore, and it was an enormous surprise to the expedition team when it was discovered living hundreds of meters high up on th slopes of the mountains. Credit: Expedition Cyclops By combining scientific techniques with the Papuan team members’ experience and knowledge of the forest, the team made a wealth of new discoveries. These included several dozens of insect species completely new to science and the rediscovery of Mayr’s honeyeater (Ptiloprora mayri), a bird lost to science since 2008 and named after famed evolutionary biologist Ernst Mayr. An extraordinary finding was an entirely new genus of ground and tree-dwelling shrimp. “We were quite shocked to discover this shrimp in the heart of the forest, because it is a remarkable departure from the typical seaside habitat for these animals,” said Dr. Leonidas-Romanos Davranoglou (a Leverhulme Trust Postdoctoral Fellow at the Oxford University Museum of Natural History), lead entomologist for the expedition. “We believe that the high level of rainfall in the Cyclops Mountains means the humidity is great enough for these creatures to live entirely on land.” The team also revealed a treasure trove of underground species, including blind spiders, blind harvestman, and a whip scorpion, all new to science, in a previously unexplored cave system. This astonishing discovery was made on one of the sacred peaks above Yongsu Sapari where the team had been given special permission to do research. People rarely tread here, and the striking cave system was chanced upon when one team member fell through a moss-covered entrance. About the Expedition: The expedition was first proposed in 2019 by James Kempton. Academics who collaborated closely on the expedition include Dr. James Kempton (University of Oxford), Dr. Leonidas-Romanos Davranoglou (Oxford University Museum of Natural History), Madeleine Foote (University of Oxford), Dr. Andrew Tilker (Re:wild, Leibniz Institute for Zoo and Wildlife Research), Dr. Attila Balázs (Mendel University) and Dr. Max Webb (Royal Holloway, University of London). Cenderawasih University (UNCEN) team members and partners include Dr. Suriani Surbakti, Gison Morib and Heron Yando. Team members and collaborators from Indonesian NGO Yayasan Pelayanan Papua Nenda (YAPPENDA) include co-founders Iain and Malcolm Kobak, and Yali Kobak, Sampari Kobak, Ezra Daniel, Ruben Penggu, Melias Heluka, Yuanis Yalak and Sili Yalak. The team obtained permits from Papua BBKSDA and BRIN. They were also given permission from the community of Yongsu Sapari to conduct research and collect specimens on their land during the expedition. This rediscovery was made possible in part by the generous support of Merton College Oxford, the Royal Geographical Society, the Scientific Exploration Society, Re:wild, Royal Holloway University, the University of Oxford, Reconyx, and private donations from Derek Williams, Chris Peri, and other generous individuals. “A Beautiful but Dangerous Land” Extremely challenging and, at times, life-threatening conditions were at the background of these discoveries. During one of the trips to the cave system, a sudden earthquake forced the team to evacuate. Dr. Davranoglou broke his arm in two places, one member contracted malaria, and another had a leech attached to his eye for a day and a half before it was finally removed at a hospital. Throughout the expedition, members were beset by biting mosquitoes and ticks, and faced constant danger from venomous snakes and spiders. Making any progress through the jungle was a slow and exhausting process, with the team sometimes having to cut paths where no humans had ever been before. Cenderawasih University (UNCEN) student and team member Gison Morib setting up one of the eighty camera traps which were placed in the Cyclops Mountains in order to capture images of Attenborough’s long-beaked echidna for the first time. Credit: Expedition Cyclops “Though some might describe the Cyclops as a “Green Hell,” I think the landscape is magical, at once enchanting and dangerous, like something out of a Tolkien book,” said Dr. Kempton. “In this environment, the camaraderie between the expedition members was fantastic, with everyone helping to keep up morale. In the evening, we exchanged stories around the fire, all the while surrounded by the hoots and peeps of frogs.” About the Cyclops Mountains: The Cyclops Mountains are one of the most inhospitable regions in the world, being rugged, steep, and dangerous, and ranging from sea level to 1,970 metres. They are dominated by two main peaks – Gunung Rara and Gunung Dafonsoro. When sailing ten leagues from the range’s northern coastline in 1768, Baron Louis de Bougainville noted “les deux cyclopes,” from which the mountains bear their name. The Cyclops Mountains are a Key Biodiversity Area, meaning that they are critical to the persistence of biodiversity and to the overall health of the planet. The community of Yongsu Sapari have lived in the region for eighteen generations and hold the land as sacred. They believe it is stewarded by a female spirit who can take the form of a tree-kangaroo. An Enduring Legacy Rediscovering the echidna is only the beginning of the expedition’s mission. Attenborough’s long-beaked echidna is the flagship animal of the Cyclops Mountains and a symbol of its extraordinary biodiversity. The team hopes that its rediscovery will help bring attention to the conservation needs of the Cyclops, and Indonesian New Guinea more generally, and they are committed to supporting long-term monitoring of the echidna. Key to this work is NGO YAPPENDA, whose mission is to protect the natural environment of Indonesian New Guinea through empowerment of Indigenous Papuans. As part of the expedition team, members of YAPPENDA helped train six students from UNCEN in biodiversity surveys and camera trapping during the expedition. Dr. Davranoglou said: “Tropical rainforests are among the most important and most threatened terrestrial ecosystems. It is our duty to support our colleagues on the frontline through exchanging knowledge, skills, and equipment.” With the team having only sorted a fraction of the material collected on the expedition, they expect that the coming months will yield even more new species. The intention is to name many of these after the Papuan members of the expedition. Besides animal specimens, the team also collected over 75 kg of rock samples for geological analysis, which was led by the expedition’s chief geologist, Max Webb, from Royal Holloway University, London. These could help answer many questions about how and when the Cyclops Mountains originally formed. The mountains are believed to have formed when an island arc in the Pacific Ocean collided with the New Guinea mainland about 10 million years ago. Combined with the biological findings, this geological work will help the team understand how the extraordinary biodiversity of the Cyclops came to be.

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